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CS MALTA I AOM “NURAGIIINO reni nani ovo re er Re dO ANTE ere enen NI_NVINOSHLINS S31YVY911 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_ more = = = ' = $ a oO. — œ _ 1 œ u len, 5 5 5 = © = E > = > = PACE = > ) > = 7 = OF KR > E = m eu m LES m a m uw = vw = u = uw ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IUVY811 LIBRARIES a PZA v = u à Zs un = < = < = < = = > 2 Si 2 è = = = a 5 = ì Ó a ; 72 3 2 e = = =, E Z E 2 >" > > = = 2 A 2 7 x È NI_NVINOSHLINS S3IUYVYAI1 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI = wo — o — v 5 5 a 5 a 5 u 2 | 2 = 3 = eS Li = E po; = a (= o = = m a = Pr = o = o Si si = e 2 = = = = pr) 2 ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3iuvug11 LIBRARIES = ie = = z te = o “oO o œ o œ = AS 2 = D E a pa sì > > =| £ =>) > F I r > Pr 2 de ers 2 = a = == la no = n - > = U z i 2 z wes o £ SNI_NVINOSHLINS, S31UVHOI LIBRARIES SMITHSONIAN INSTITUTION, NOILNLILSNI È PF K È È 8 5, N Vs è i: È ip È ww: = > = > = È 3 2. 7 ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS saruvug11_L'BRARIES 811 LIBRARIES SMITHSONIAN tu zZ u 2 2 ‘ pei LS sa œ 3 = a cy DP = < = = È m 4; = ao _ o = 3 = a = = al 9 x 2 = NI _NWINOSHLIWS zei Iuvyal 1 LIB RARI ESSSMIRSONIAN INSTITUTION | = = 5 o N, © \ > N = be) = Gy, 5 > NAE > = GLE = pes NF E È WA 2 = ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS SIIUVUAIT AIWIAIOCQUTIIIAIC cqupruwuaitt (22) un | Z = < 5 E = < = = “È “i = 53 = == le) NO E & © = 9 a D WA 77] 0, [2] T \ le) TE AND 2 TE (©) = ~ 2 E NN 2, 5 Z., = 3 = NY > = > 3 u a z ” Tj NI NYINOSHLINS S3IYVYG!IT LIBRARIES SMITHSONIAN INSTITUTION | = = 2 zZ a 3 Ww 3 7) u us u = i =, © a = Zi o = a) 3 2 = = ka < È 3 = N x Land & = x - =, EN SS co =. ai re] 7 = RR A 5 = 5 - È z a = = = + 7 ES SMITHSONIAN INSTITUTION NOILNLILSNI NYINOSHLIWS SALUS LIBRARIES =a = a rm z IE « 9 0 2 pen NE © w ME ct aS = 5 NY = = FI AD 3 > >| € K = > 5 E ty 4 be > N + - 9 > È > = > È y = sia Fr A € oO m [2 m n m : i | Wi I ci 8 u » 1 1 al aN 7 G | " ri RTL See ee (2 u D. | 7 sE nn, DT DE Te ’ Fe Er ME ROUE 1. Des È DAT Fr = by i . ( 0 i = DI oral CAGES i n 7 I m, ® 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 È „al | i A gi u f i ai u 4 A oe (ia so | riko ei 7 b { | | er ar! ito? un Ay; wes er 1 | À fas ” i | NI | ' dl iat Ab L u ol nn Î k ehi 1 { i i la nai gee rol [a ı A ! N inns tn ! il INTEL mai { IE 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 PAUL | MAY 18197 = / NCIBRARIE 2 4 \ NEDERLANDSE ENTOMOLOGISCHE VERENIGING BESTUUR (BOARD) Voorzitter (Chairman) . . . + G. Barendrecht Vice-Voorzitter (Vice- President): a ts) ANT. Wiebes Secretaris (Secretary). . » - . ++ + W. Hellinga “Addresc 20. 4. « nn. … Weesperzijde 23 Il, Amsterdam-1005. Penningmeester (Treasurer). MAIER WATMKabos Address . . . un... Van Baerlestraat 26 I, Amsterdam-1007. Bibliothecaris Arihrariany I EVEN. RELITS Address. > 2: 2 « +» « +» . + + Zeeburgerdijk 21, Amsterdam-1006. Leden (Members) . . . . . . . . . G. van Rossem, A. F. H. 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The Journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. It appears in separate issues, forming an annual volume of 350—400 pages. Subscription rate: D.Fl. 62.50 per volume. MONOGRAFIEEN VAN DE NEDERLANDSE ENTOMOLOGISCHE WERENIGING Redactie (Editorial Board) and address as for Tijdschrift voor Entomologie. The Monographs are intended for the publication of larger entomological papers on a single subject and will appear irregularly. The following Monographs have been published: Hors série: F. T. Valck Lucassen et al., 1961. — Monographie du genre Lomaptera Gory & Percheron (Coleoptera, Cetoniidae), 299 pages, 739 figs, 2 pl., map. D.FI. 60.—. No. 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. SYSTEMATICS AND GEOGRAPHIC HISTORY OF THE GENUS PYRGUS IN THE PALAEARCTIC REGION (LEPIDOPTERA, HESPERIIDAE) by 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, ...” (c) Sichel (1962) concluded from morphologic, chorologic and palaeo-zoogeographic 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. (c) Female genitalia: Fig. 3. 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 28 1 mm db 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 35 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 WE en Ee serratulae-t - type Sr = 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] : en 14 . î 73 12 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 59 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 y 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 (c) scandinavicus. — The scandinavicus type of genitalia is found as well in Scan- 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 AE Vn | Su Ay à om dl PY RS 9 dI N Map 1 - Pyrgus medal re nn au | FR = - 3 de (i) Hi \ p WC, È g 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 Eh RS 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 4 LAN a) gar i Map 7 =“ | N \ NND Superspecies Pyrgus alpinus N SAT = È \ aq Pyrgus alpinus | r & 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 SS 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 di ‘I FO à In i Je {N LI in ut Nd 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. —, 1938. — Ueber das Vorkommen von Hesperia carthami Hbn. in Norddeutschland, nebst Bemerkungen über Verbreitung und Formen dieser Art. — Stett. ent. Zeit. 99: 236—246. —, 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. 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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 . u ® u ) _ * | Se Sele ek dope aT ware Sm = sae ‘6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 1, 1972 PLAAT 1 16 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 We 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 No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.Fl. 70.—. No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. D.Fl. 25.—. No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. D.FI. 100.—. 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The Monographs are intended for the publication of larger entomological papers on a single subject and will appear irregularly. The following Monographs have been published: Hors série: F. T. Valck Lucassen et al., 1961. — Monographie du genre Lomaptera Gory & Percheron (Coleoptera, Cetoniidae), 299 pages, 739 figs, 2 pl., map. DFI. 60.—. No. 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.FI. 30.—. SOME OBSERVATIONS ON THE BEHAVIOUR OF FEMALES OF PHILANTHUS TRIANGULUM (F.) (HYMENOPTERA, SPHECIDAE) ') by 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 WM VVM VL TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972 À 13 12.” 18 17? -19 14 16 15 August 21. 23 25 UA 3775 ed a EE CS CE u) a RG B1 B1 CYR S5——S5 G7 G7 BY ? YW 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 1 outer closure A Pr + main burrow cocoon x LU 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 © D = D co 5 @ © 5 4 = © D O2 [=] © © O © 20 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 50 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 o = © 5 $ 5 15 A 2 o 10 — o mR © 2 £ 0 € = 60 120 240 360 min 60 o S 2 40 ~ 5 © B 7) Ro) o bo o © 2 20 E 5 = 0 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 — a number of units of watching — o (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 12 œ number of g/day o > number of d/day N 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. ye ee u i zl LÉ ph te Lun i di > u RT peat ere ape Fr LUTTE aye! prete are L “À ne ur vivi ni hp or P hairy? OF a — namvech li =" M Fe È Mei Lot è pe pal Gu 499 A mare ai PI sel cd vient u Saisies A | PA 5.10 ds . | L NT a 4 re PR sr tnt ae "ef BEND . ze è x = hi Auivesr © per J mth, ee n ik Ù | : Re DDL dite Comes Le CE sp : Fu = Ce in V pit doh) bij y cs set e A u u saf É > (eng 8 2 u Dr pp! © on 1 » a l 1 - «ni il vr ai met sen alé n NOTES ON THE BEHAVIOUR OF MALES OF PHILANTHUS TRIANGULUM (F.) (HYMENOPTERA, SPHECIDAE) by 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 pozeorpur are (ca) wnpnSuviag suqqurpidd JO sojew UAAIS JO SUOTJEJS JELIONIII9) PUE "}OYIS snyoswwyN snqny ‘7 winyAduas snwAyL ‘7 Nog eat YI ‘INN snuje eynbues4 ays Buipeay uol}2}S |2110}1119} ajoy Buideajs Ad dA 9 ‘ Says FUIP33F sajoy surdaays € ‘(aoueig) opnogeN je vore Ápms ayy, ‘I ‘Sig 143 4 le) R. T. SIMON & E. P. R. POORTER : Behaviour of Philanthus 6961 WOE Pur y97 aunf uo (M) (‘4) UNINGUPIAL SNGIUP IGT oew e JO SONIAIDY ‘7 ‘SIA UO!}e3S [8110311187 UO 9 Mm apisui 9 ‘uado ajoy apisjno 9 | says Bulpaajy uo 9 apisui 9 ‘paso|jo a|oy ‘wd g v rd zl "We OL 144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 2, 1972 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 Me | 69 | number o observation 80 males present at the same time 60 0 d 1d 40 26 36 20 N N > = é LE) ili EVNYNYUNUNWMAAN, 10 a.m. 12 2 4 6 p.m. Fig. 3. Relative frequencies of numbers of males at territorial stations in the course of the day 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 WIW+Y* |W+Y*| L G BA 5/o|4/o |3/2| 3/3 = 3/0 Ea 1/0 [1/0 [1/4 n = S | ,o Li zn EAN = uler|sa aN [EN = Fl] N n au EN n S avi zn n Na QUI GS = n a chat ti LÌ * F S n n am n n YU ASIN Nun ol un KN TKN Wu mE PIENI oN] aN nv} ou *| + U ER HU | nd u an n LA dele bilan = D as an Ci LI YI S LES Hn IS IN n EN > Le] & = (©) + SS = AN de NES o|dn 0 LI = © Li [BAN | OAN NS [ES [BN AO = n 7 WG |WG WG WG | WG WG pEr a BIG GW |GW+RW 2 RY RY [RY |RY RY RY RY RY |RY |RY di ARE PG |PG |PG PG PG PG BR |BR |BR BR | BR+B* B* |B* |B* valse rl 20 ele VATIIATATA B* n A B* B* B* Be B* B*+B* |B* | Be |B* |B* Be B* Be B* B* B* Be |B* |B* oie celal 2, P P P 3/1| 3/2 |3/1| 3/1 |2/1 [1/1 |1/0]1/0 1/1 1/1| 3/1 |3/0| 3/4 |2/1|1/1 Li 1/0 ta EEE ra are pe are ro DADACATOLA u0T9638 TETIOFTII9I 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 MXX ENE OT GTS TONI or AR 24 104 MXXX D TOR ee 3 XX ONO ON RON GLP a Oy Oe OO) 2.) x (eee ORTA O I ON OE OT rile GENO SA 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. be Ghose On | x u prc = (= van 4 BR TEN a (me Salone hasta Balt i a dio dites re : can Fun ge sw = EME) ’ Ait =o =a @ MIS PARMI Ì i Pag) mer n © Vi Pf att. ae ah hee A ‘ = Mii a il yo ed pae e AP At MORE LT IP cina a © hs a Ere (pe n Te mimi wah > it ER veh etn ne 5 une owl Al Ni, x lai, @ 5 nq p= ing ae wu a i | df Tide ace Pao) ca UN ln * Md ety * & » ox . ks 140 Oul wees ns Mes, AB sl ice A u Ir Ye A Cling ZA, FE 7 hè creto ad { [FU ti a Un LULA wgn Bl; 5 mi i _ “Ag @ | | 8 i 12 CRE (re. | gin at Je ie a ai rer Gi Goal? rile — ila ria n À id 7 a i i ig € vi Ta MA à lisse one eef ei MI din (1 er i Ana a oe” u ee ee n recai | Ava bal rus die Fehr î TT : ‘i nal) oe 0001 sl ni j > u e © No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff. — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.Fl. 70.—. No. 4. F. Willemse. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. D.FI. 25.—. No. 5. C. A. W. Jeekel. — Nomenclator generum et familiarum Diplopodorum. 412 pp. D.Fl. 100.—. ENTOMOLOGISCHE BERICHTEN The Journal serves the publication of short papers, faunistic notes, reports of the meetings, etc. It appears monthly in issues of 16—24 pages, forming one volume annually. Redacteur (Editor) ere BJ Lempke Address Dares . . . . . Oude IJsselstraat 12 III, Amsterdam-1010. Subscription rate: D.Fl. 40.— per volume. 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Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. CONTRIBUTION TO THE KNOWLEDGE OF ORIENTAL PSENULUS (HYMENOPTERA, SPHECIDAE, PSENINI) by J. P. VAN LITH Rotterdam ABSTRACT The following new species and subspecies are described and illustrated: P. biszcatus, W Pakistan; suifuensis, China; parvidentatus, Formosa; nasicornis, Celebes; bengalensis, India; nietneri, Ceylon; pallidicollis, Borneo; neptunus, Celebes; pseudajax holtmanni, Culion and Busuanga (Philippine Is.) ; interstitialis salomonensis, Guadalcanal, Malaita, Russell Is., Kolombangara, Vella Lavella, Bougain- ville (Solomon Is.); segrex, Celebes; chariis, Choiseul (Solomon Is.); arancarius, New Guinea; lamprus, Celebes; annamensis, Vietnam; hoozanius, Formosa; ceylonicus, Ceylon. New records of known species are given. From various regions, particularly continental southeast Asia, Celebes and the West Pacific islands, interesting material has been received during the last few years. It con- tained many new species, or the opposite sex of forms described earlier after only one sex. This enabled me to obtain a better insight into the division of the genus into species- groups and their relationships. The more our knowledge of the species of Psenulus increases, the stronger become grounds for the erection of a number of new subgenera. The only subgenus published so far is Eopsenulus Gussakovskij, 1934, for iwatai Guss., whilst the genus Nzpponopsen Yasumatsu, 1938, was erected for anomoneurae Yas. (syn. Psenulus mandibularis Tsune- ki, 1959). The new subgenera should be based mostly on the already distinguished species groups, or on combinations thereof. I hope to discuss this question in a future paper, after a study of the African and New World Psenulus. I express my gratitude to the institutions and entomologists who have sent me material. Their names are listed below together with the abbreviations as used in the present paper. BISH — Bernice P. Bishop Museum, Honolulu BM — British Museum (Natural History), London BPIM — Bureau of Plant Industry, Manila CAS — California Academy of Sciences, San Francisco CNC — Entomological Research Institute of the Canadian Department of Agriculture, Ottawa HUA — Hyogo University of Agriculture, Sasayama Coll. Krombein — Dr. Karl V. Krombein, Washington, D.C. MCG — Museo Civico di Storia Naturale, Genoa ML — Rijksmuseum van Natuurlijke Historie, Leiden OUM — University Museum, Oxford Coll. Pagden — Mr. H. T. Pagden, Penang, Malaya 153 154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Coll. Roche — Mr. C. G. Roche, Singapore Coll. Townes — Dr. Henry K. Townes, Ann Arbor, U.S.A. UDE — University of Delhi, Department of Entomology, Delhi USNM — National Museum of Natural History, Smithsonian Institution, Washington, D.C. (formerly United States National Museum) ZMB — Museum für Naturkunde der Humboldt-Universität, Berlin Group of Psenulus fuscipennis Psenulus bisicatus spec. nov. Female. — Black; palpi light brown; last 7 or 8 segments of antennae reddish-brown below; legs dark brown; foreside of fore tibiae, anterior tarsi, mid tarsi and ends of tarsal segments of hind legs light brown; base of hind tibiae slightly paler than rest of hind tibiae. Apical spines of fore legs light brown, of mid and hind legs whitish. Veins of wings dark brown. Interantennal carina raised and very broad, nearly twice as broad as diameter of ocelli, lozenge-shaped, excavate and with very fine sculpture. The lateral carinae of this broad- ened part do not fuse below, but are parallel for a short distance, ending in transverse carina below antennae. This carina arched in frontal view, with an angle of slightly more than 90 degrees. Below this carina a smooth and shining triangle with a few large punc- tures, remainder of face and clypeus with dense and fine puncturation, without distinct interspaces. Anterior margin of clypeus with two long triangular teeth; distance between these teeth as long as depth of the rounded emargination (Fig. 1). Anterior margin of labrum with two small shining teeth. Frons distinctly and densely punctate, interspaces shining, punctures with tendency to oblique striation. Vertex sparsely punctate, smooth. Antennae short, clavate, third segment about one and a half times as long as broad at apex, segments 8—11 about as long as broad, 12th segment about one and a half times as long as broad at base. Prescutal sutures visible only on anterior half of scutum. Surface of scutum shining, finely punctate, interspaces on the whole a few times larger than punctures. Scutellum very shining, almost impunctate. Enclosed area of propodeum depressed, narrow, with distinct longitudinal oblique carinae, median area broader than long, with median longi- tudinal carina. Upper part of posterior side of propodeum smooth and shining, lower third part with minute, hair-bearing punctures. Median longitudinal sulcus of propodeum broad and shining, ending before apex, with a few transverse carinae divided over the whole length. Sides of propodeum with fine oblique striation. Mesopleura with sparse and fine puncturation, a few indistinct transverse striae. Anterior oblique suture and its widened upper part with distinct transverse carinae. Legs normal. First recurrent vein of fore wings ending just in second submarginal cell, second recurrent vein ending well in third submarginal cell (distance slightly more than one fourth of base of third cell). Petiole not reaching as far as apex of hind femur, dorsally with shallow groove which is widened at base; rounded below. Gaster finely punctate. Lateral carinae of pygidial area about one third of length of last tergite in normal situation, apex rounded, surface dull (Fig. 2). Second gastral sternite with a distinct semi-elliptic depression. Face below transverse carina silvery pubescent, pubescence of clypeus appressed only J. P. van LITH: Oriental Psenulus 155 laterally. Pubescence of thorax and gaster whitish and of normal density, hind margin of fourth and fifth gastral sternites with a fringe of long white hairs. Pygidial area with fine and dense, long pubescence. Epicnemial areas below with a large round patch of fine, not very dense pubescence. Back of propodeum except narrow upper margin with fine, long pubescence. Length about 6,5 mm. Male. — Resembling female, with following exceptions. Third and following seg- ments of antennae reddish-brown below. Apex of gaster and apical spine brown. Clypeal teeth much shorter than those of female, emargination more rectangular- triangular. Interantennal carina about one and a half times as broad as ocelli or, at most, as broad as first antennal segment, in the female this carina about one and a half times as broad as first antennal segment. Upper part of frons and vertex with oblique striation. Antennal segments 3—11 at least one and a half times as long as broad at apex, 13th segment about twice as long as broad at base, segments 3—8 with linear, brown and shining tyloidea, on ninth segment reduced to a point. Posterior surface of propodeum with coarse carination, but almost smooth behind enclosed area of propodeum. Second recurrent vein of fore wings ending more close to base of third submarginal cell. Length about 6 mm. W Pakistan: 2 9, holotype and paratype, Quetta, Aug., 1903, 1 3, allotype, Aug. 1903, 2 d', paratypes, June and Aug., 1903, coll. Col. C. G. Nurse (BM, 1920—72). P. bisicatus is closely related to P. atratus (Fabr.), from which it differs by the broader interantennal carina and the smooth posterior surface of the propodeum. It is easily distinguished from P. meridionalis de Beaumont, which also has a broad interantennal carina, by the larger extent of the smooth part of the propodeum, the shining and very finely punctate vertex, and the finely punctate scutum, and from P. pan de Beaumont, described from Turkey, by the longer antennal segments of the male and the longer clypeal teeth of the female. It differs from P. laevis Gussakovskij, which according to the description also has a sparsely punctate mesonotum, by the long clypeal teeth and by the colour of the legs. Group of Psenulus suifuensis Psenulus suifuensis spec. nov. Female. — Black; underside of last four antennal segments very dark brown; tegulae, fore and mid tarsi and apices of hind tarsi dark brown. Veins of wings black. Clypeal margin protruding medianly, distinctly bidentate (Fig. 3), disk densely punctate. Face above clypeus also densely punctate, medianly longitudinally striato- punctate. Frontal carina raised and ovally broadened between antennae (Fig. 5), deeply excavate. No transverse carina below antennae. Frons densely punctate. Vertex behind ocelli transversely striate. Sloping posterior surface of vertex laterally almost impunctate and shining, tempora very finely obliquely striate, occipital carina not ending in hypostomal carina (Fig. 4). Third antennal segment about two and a half times as long as broad at apex, fourth and fifth segments about one and a half, following segments about one and a third times as long as broad at apex, last segment about one and three quarters times as long as broad at base. Mandibles rather broad. 156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 8 Fig. 1—8. Species of South Asiatic Psenulus. 1—2, head and pygidial area of female of P. bisicatus sp. nov.; 3—5, head in frontal, ventral and dorsal view of female of P. swifwensis sp. nov.; 6—8, head, interantennal carina in dorsal view, pygidial area of female of P. bicinctus Turner J. P. van LITH: Oriental Psenulus 157 Scutum irregularly punctate, median part with large impunctate shining areas. Poste- rior margin of scutum striato-punctate. Prescutal sutures about as long as anterior third of scutum. Scutellum with elongate punctures. Metanotum indistinctly punctate. Enclosed area of propodeum with oblique longitudinal carinae, area between median two carinae irregularly carinate. Posterior surface of propodeum shining, with a few fine, hair- bearing, punctures. Sides of propodeum with indistinct sculpture. Mesopleura and mesosternum with fine punctures. Legs normal, outer side of hind tibiae with one small thorn, at a distance of about one-fourth of total length from apex. First recurrent vein of fore wings ending in second submarginal cell, which is pentagonal; second recurrent vein ending in third submarginal cell. Apex of petiole reaching about as far as apex of femora. Underside of petiole raised into an indistinct keel. Sides with shallow groove and fine, irregular, punctures. Base of petiole dorsally with V-shaped carina, reaching beyond middle, apical part with shallow, broad, groove. Gaster with fine punctures, hind margins of tergites almost impunctate, pygidial area very weakly indicated. Second gastral sternite with ill-defined, almost impunctate, semi-elliptic depression, rest of gastral sternites finely punctate. Face with appressed silvery pubescence and long, erect, whitish hairs. Ventral side of thorax with dense, whitish, short pubescence and also with long hairs. Epicnemial areas below with circular patch of yellowish-grey pubescence. Fourth and fifth gastral sternites with dense fringe of short, whitish, hairs. Sixth sternite densely whitish-pubescent. Rest of body greyish-pubescent. Length about 9 mm. Male unknown. China: 1 © (holotype), Suifu, Szechuen, April-May, 1928, coll. D.C. Graham (USNM). This is one of the few species of Psenulus in which the occipital carina does not end in the hypostomal carina, while the transverse carina below the antennae is lacking. Group of Psenulus rufobalteatus Psenulus rufobalteatus Cameron 1904, Cameron, Ann. Mag. Nat. Hist. (7) 13: 219, ® (Psen rufo-balteata; Assam). 1962, Van Lith, Zool. Verh. Leiden 52: 111. 1965, Van Lith, Zool. Verh. Leiden 73: 59. Original description: “Black; the apex of the second abdominal segment and the whole of the third rufous; the fifth and following joints of the antennae testaceous beneath. Legs thickly covered with white hair; the spurs pale rufous. Wings clear hyaline; the first cubital cellule in front is half the length of the second, the first recurrent nervure is received very near the first transverse cubital, the second at twice the distance from the second transverse cubital. 9 . Length 5 mm. Hab. Khasia Hills. Coll. Rothney. Antennae stout, thickened towards the apex; the scape beneath sparsely covered with long black hair. Face and clypeus thickly covered with silvery pubescence; the front and vertex almost bare, sparsely punctured; the eyes almost parellel; the ocelli in pits; the 158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 front with a shallow central furrow; antennal tubercle large, the apex triangular, its sides distinctly margined, the middle depressed; below this is a larger one, roundly incised at the apex, the sides rounded. Pro- and mesothorax shining, sparsely covered with white hair; the mesonotum distinctly but not closely punctured; the scutellums with a few fine punctures. Area on the base of metanotum narrow, elongate, and marked with stout striae; the central furrow is wide and deep, becoming slightly wider towards the apex, and marked with a few stout striae; on either side of it at the apex is a large leaf-like expansion, its apex transverse, on the outer side covered with long hair; at its base is a small rounded projection. Pro- and mesopleurae finely and sparsely punctured; below the mesopleural tubercles is a wide, deep, slightly oblique furrow, marked with some trans- verse keels; there is a smooth furrow near the base of the metapleurae; the apex of the latter is rugose and is marked with some transverse keels. Comes near to P. rufiventris, but is quite distinct therefrom.” The holotype, which is in the type-collection of the Oxford University Museum, is badly damaged. The gaster is partly missing, the left antenna is broken off after the third segment and there is a hole in the sixth segment of the right antenna. A redescrip- tion based on this specimen is given below. No further material has been collected. Female. — Head black, palpi testaceous, middle part of mandibles dark reddish, under- side of fourth and fifth antennal segments brown, remainder of underside light brown. Upper side of antennae somewhat darker. Thorax completely black, tegulae and legs dark brown. Tibial apical spurs yellowish-white. Veins of wings very dark brown. Petiole black, first gastral tergite dark brown with somewhat transparent margin, second tergite and sternite pale red, base of third tergite and, according to the original description, also the rest of the gaster black. Median part of anterior margin of clypeus with two triangular teeth, margin much depressed. Anterior margin of labrum laterally with a long tooth. Frontal carina fine but distinct, raised interantennal part much broadened, lozenge-shaped, below the antennae this carina ends in a transverse carina, which, in frontal view, has the shape of a broad inverted V. Between this transverse carina and the clypeus, a large, distinct, circular depression. Face below transverse carina densely punctate. Frons and vertex shining, below ocelli finely but distinctly punctate, vertex sparsely punctate, with very slight tendency to oblique striation. Antennae rather short, clavate, 10th and 11th segment almost as long as broad at apex, last segment nearly twice as long as broad at base. Scutum shining, finely but distinctly punctate, interspaces generally twice or four times as large as punctures. Prescutal sutures and parapsidal sutures normal, distinct. Scutellum and metanotum sparsely punctate. Enclosed area of propodeum depressed, longitudinal carinae sharp, median pentagonal area with irregular median longitudinal carina. Posterior surface of propodeum shining, smooth behind enclosed area, lower part finely punctate. Sides of propodeum dorso-laterally with coarse carination, forming a row of large, almost square, depressions. First recurrent vein of fore wings ending in second submarginal cell, second recurrent vein ending well in third submarginal cell. Petiole rounded below, dorsally with shallow longitudinal groove, laterally with low longitudinal ridge, above this, on basal half, a much sharper carina. Pubescence long and whitish, dense on thorax. Epicnemial areas below with large and round patch of silvery pubescence. J. P. van LITH: Oriental Psenulus 159 North India: 1 9, Assam, Khasia (OUM). This specimen was labelled “type” by Cameron. P. rufobalteatus is closely related to P. bicinctus Turner, from Shillong (Assam). In the latter species also the last gastral segments are red. The circular depression below the transverse carina of the face is less deep — in the Palaearctic P. concolor (Dahlbom) this character is not constant! — the antennae are dark brown below instead of light brown. The following error in my 1965 paper should be corrected: in the description of Psen rufoannulatus, the 18th sentence on page 59 should read: “a. I could not find any other specimen of P. rufoannulatus” etc. (instead of “P. rufobalteatus”). Psenulus bicinctus Turner 1912, Turner. Ann. Mag. Nat. Hist. 8 (10): 363-364, 2 (Assam). Redescription of female. — Head and thorax black; apex of mandibles dark reddish, underside of antennae brownish, palpi and tegulae dark brown. Legs dark brown. Veins of wings black, stigma brown. Petiole including ventral plate, first, third, and fourth tergites, except hind and lateral margins, black. Rest of gaster, including second, fifth and sixth sternites red. Disk of clypeus slightly convex, depressed anterior margin bidentate. Clypeus and face coriaceus. Interantennal carina raised, broadened and excavate between antennae (Fig. 7), narrowing into transverse carina below antennae (Fig. 6). Frons densely and finely punctate, vertex more widely punctate. Interocellar area very weakly rugose. Antennae somewhat clavate, third segment nearly two and a half times as long as broad at apex, segments 9—11 slightly longer than broad at apex and last segment about one and three quarters times as long as broad at base. Scutum distinctly punctate, interspaces on disk generally a few times as large as diam- eter of punctures, denser near prescutal sutures, which are distinct only on anterior half of scutum. Parapsidal sutures fine, connected with hind margin of scutum by a deeper groove. Scutellum and metanotum finely punctate. Enclosed area of propodeum narrow, strongly foveolate and sometimes separated from metanotum by a triangular rugose area. Longitudinal sulcus on back of propodeum broad, central part deepest, above and below this deeper part a few transverse carinae. Propodeum on dorsal part shining, posterior surface with fine punctures, separated from the sides, which have a reticulate carination, by a foveolate groove. Mesopleura and ventral side of thorax finely punctate. Anterior plate of mesepisternum finely punctate. Anterior oblique suture coarsely foveolate, widened upper part with a few transverse carinae. Legs normal. First recurrent vein of fore wings ending in first, second recurrent vein in third submarginal cell. Only apical third of petiole above weakly sulcate, sides somewhat keeled, ventral side rounded. Gaster finely and widely punctate. Pygidial area (Fig. 8) elongate-triangular, slightly narrowed towards apex, finely punctate. Base of second gastral sternite with indistinct, semi-elliptic depression, not well-defined. Face with silvery appressed pubescence, also with long erect greyish hairs. Vertex and thorax with long greyish hairs. Underside of thorax densely whitish-pubescent. Epi- cnemial areas below with a patch of dense pale pubescence. Gaster with short yellowish pubescence, longer and denser at apex, hind margin of fourth and fifth gastral sternites 160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 with short and dense pale pubescence, sixth sternite with a large patch of dense golden- brown pubescence. Length about 8 mm. First description of male: Pronotal tubercles slightly paler, dark brown. Veins of wings brown. First tergite only on basal half black and of third tergite only posterior part black. Apical spine dark brown. Clypeal disk more strongly convex. Interantennal carina broadened and excavate, as in female and ending below in a transverse carina. Frons densely but finely punctate, vertex shining, with a few small and some larger punctures, less densely punctate than in female. Antennae slender, third segment slightly longer than twice, following segments about twice as long as broad at apex, last segment nearly two and a half times as long as broad at base. Antennal segments 3—11 with long, somewhat oblique tyloidea, on last segments gradually diminishing, on 12th segment indistinct. Prescutal sutures not distinct. Propodeum as in female, but apical third of back of propodeum with coarse reticulate carination. First recurrent vein of fore wings interstitial. Petiole dorsally with distinct longitudinal groove, edged laterally, sides with narrow groove, ventral side of petiole rounded. Second gastral sternite without semi-elliptic depression. Face with dense and appressed, silvery pubescence and also with long, erect, greyish hairs. Length about 6,5 mm. Assam: Turner recorded four females of which I have seen two specimens: Shillong, July, 1903, coll. R. Turner 1905—125 (BM). New record from Assam: 1 9, Upper Shillong, 5000 ft, 2 May, 1962, coll. V. Gupta (UDE). Sikkim: 1 4 (allotype), Tsuntang, 6000 ft, 24 April, 1924, coll. Major W. G. Hing- ston, Everest Exp. Brit. Mus. 1924—386 (BM). Nepal: 1 9, Dunche, 28 km N of Trisuli (Nawakot), 1950 m, 7—12 Nov., 1965, coll. L. W. Quate (BISH). This female is darker than those from Assam. Mandibles and antennae almost completely black; femora and tibiae black, instead of brown; fourth tergite completely and basal half of fifth gastral tergite, black. Narrow hind margin of third, fourth and fifth sternites reddish. Interocellar area distinctly punctate. Below transverse carina a circular depression, more distinct than in the females from Assam. Parapsidal sutures not connected with posterior groove of scutum. Groove on posterior dorsal side of petiole about half as long as length of petiole. First recurrent vein of fore wings ending in second submarginal cell. I am inclined to consider P. bicinctus as a subspecies of P. rufobalteatus, if not as identical, but in the original description Cameron stated that the gaster of the latter form is black and only the apex of the second abdominal segment and the whole of the third are rufous. Unfortunately the greater part of the third and following segments of the type of P. rufobalteatus are missing. Cameron did not mention the abdomen being damaged. J. P. van LITH: Oriental Psenulus 161 Group of Psenulus scutatus Psenulus scutatus scutatus (Rohwer) 1921, Rohwer, Phil. JI. Sc. 18 (3): 310-312, ® and 4 (Diodontus scutatus; Luzon). 1923, Rohwer, Phil. Jl. Sc. 22 (6): 596. 1962, Van Lith, Zool. Verh. Leiden 52: 23-27 (Psenulus scutatus scutatus; Luzon, Samar, Negros). New records from Luzon: 1 9 and 2 &, Atimonan, prov. Quezon, 1—8 July, 1908, coll. Micholitz (ZMB); 1 2, Montalban, prov. Rizal, 25 March, 1954, coll. P. Feliciono (BPIM); 1 9, 25 March, 1965 and 1 g', 3 March, 1965, Mt. Montalban, prov. Rizal, Wa-wa Dam, 150—200 m, coll. L. M. Torrevillas (BISH). Complementary descriptions are given here. Female. — Third antennal segment about two and a half times as long as broad at apex, segments 9—11 shorter than broad at apex, last segment nearly one and a half times as long as broad at base. Longitudinal sulcus on posterior side of propodeum narrow, near apex ending in a short transverse carina. Anterior oblique suture almost linear, widened upper part smooth. Second gastral sternite with usual basal depression, no well-defined semi-elliptic depression. Male. — In the male from Mt. Montalban the whole narrow area of the scutum between the median scutal lines is also black. The subalar area of the fore wings has a yellow spot. All coxae yellow. Antennae moniliform, dorsal part black, underside yellow, third segment about twice as long as broad at apex, segments 4—8 about one and a half times as long as broadest part, segments 9—11 gradually shorter, 12th segment about as long as broad and 13th segment about twice as long as broadest part. Third segment somewhat flattened, segments 4-12 strongly convex on ventral side. Anterior oblique suture very strongly foveolate, foveoles open to the rear, widened upper area with one transverse carina. Mesosternum with short transverse carinae on either side of median carina. Face with silvery appressed pubescence and also with long erect hairs; mesoster- num not densely pubescent. The extent of the black marking of the thorax, especially of the scutum, is considerably larger in the male than in the female. This great colour dimorphism is unusual in the genus Psenulus. The marking of the scutum is variable in the females and it may be reduced from a large median spot on the scutum, extending outwards over the prescutal sutures, to two separated, small and square, black spots between the prescutal sutures, as in the first mentioned female from Montalban, or even the scutum may be completely yellow, as in the female from Mt. Montalban, Wa-wa Dam. At first glance this latter specimen might be confused with P. baltazarae luteus (Van Lith, 1962) but the clypeal margins are different. Psenulus baltazarae baltazarae Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 31-32, 2 (Luzon). New record: 1 9, Luzon, Mt. Montalban, Rizal, Wa-wa Dam, 150—200 m, 10 March, 1965, coll. H. M. Torrevillas (BISH). In this female the marks on the outer side of the prescutal sutures are reduced to an irregular, vague, pale brown spot over the parapsidal sutures, slightly longer than one- fourth of the length of the scutum. 162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Group of Psenulus quadridentatus Psenulus quadridentatus Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 37-38, ® and & (Malaya). New record: 1 9, Vietnam, Fyan, 1200 m, 11 July—9 Aug., 1961, coll. N. R. Spen- cer (BISH). This specimen agrees fully with the description of the Malayan specimens. The semi- elliptic depression of the second gastral sternite is not well-defined. The hind margins of the fourth and fifth gastral sternites are covered with dense and short pale pubescence. Sixth gastral sternite with longer, backward-directed, golden-brown pubescence. Psenulus separatus Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 38, 9 (Java). 1967, Van Lith, Ent. Ber. 27: 19-20, 2 (Borneo). New records: Borneo, 1 9, Sarawak, Mt Dulit, 4000 ft, Moss forest, undergrowth, 15 Oct., 1932, Oxford Univ. Exp., coll. B. M. Hobby and A. W. Moore (BM 1933— 254). I have compared this female with the holotype. Mandibles, pronotal tubercles, tegulae and legs are somewhat more reddish yellow. Fore coxae completely brown (partly yellowish-white in the female from Java), mid and hind coxae reddish with brown base (in the holotype hind coxae completely yellowish-white). The scutellar suture shows only one median, indistinct, short longitudinal carina; in the holotype there are also a few indistinct lateral carinae. Pygidial area as in the female earlier recorded from Borneo, i.e. slightly broader than in the female from Java. Length nearly 9 mm. The original description should be supplemented as follows: axillae rather flat and appearing almost rectangular, lateral parts of scutellum little depressed. Hind margin of scutellum laterally with rectangular, densely punctate patch, covered with dense yellowish- grey pubescence. Apex of mid tibiae with a number of strong thorns. Psenulus parvidentatus spec. nov. Female. — Head and thorax black; mandibles largely dark reddish, palpi testaceous, apex of scape and underside of flagellum reddish-brown, pronotal tubercles dark brown, tegulae yellowish-red. Trochanters, apices of femora, fore tibiae, fore side of mid tibiae, base of hind tibiae and fore and mid tarsi yellowish-brown. Veins of wings dark brown. Petiole yellowish, except extreme apex and ventral plate, which are brown. Gaster red- dish, first gastral tergite dorsally largely blackish-brown. Disk of clypeus with fine punctures, apical margin depressed and with two small teeth, distance between these teeth about one-eighth of total distance between the eyes (Fig. 9). Interantennal carina between the eyes raised, much broadened (Fig. 10) and concave; below the antennae the interantennal carina ends in a short, arched, transverse carina (Fig. 9). Frons and vertex smooth and shining, with extremely fine and wide- spread puncturation. Antennae somewhat clavate, third segment nearly two and a half J. P. van LitH: Oriental Psenulus 163 times as long as broad at apex, segments 9—11 broader than long, last segment about one and a half times as long as broad at base. Scutum with fine widespread puncturation and a few larger punctures. Prescutal sutures distinct on anterior two-thirds, vague on posterior third of scutum. Scutellum and metanotum with widespread punctures. Enclosed area of propodeum short, triangular, no median longitudinal carina. Back of propodeum smooth, upper part shining, without hair-bearing punctures. Median longitudinal sulcus narrow. Sides of propodeum almost smooth, edge between back and sides with some irregular carination. Anterior oblique suture coarsely foveolate, widened upper part with some transverse carinae. Hypo-epimeral area distinct. Legs normal, apex of mid tibiae with four red spines. Second submarginal cell of fore wings normal; first recurrent vein ending in first submarginal cell, almost interstitial, second recurrent vein ending in third submarginal cell, distance from second submarginal cell about one-fourth of base of third. Petiole cylindrical. Pygidial area long and narrow, lateral carinae distinct (Fig. 11). Second gastral sternite with indistinct semi-elliptic depression. Pubescence of face silvery, leaving sculpture visible, appressed and also with long greyish hairs. Depression of interantennal carina laterally with a row of long hairs. Back of propodeum densely greyish-white pubescent. Rest of body also with greyish-white pubescence, denser on mesosternum. Epicnemia below with a round patch of golden- brown pubescence. Margin of fourth and fifth gastral sternites with dense, short, greyish pubescence. Length about 7 mm. Male unknown. Formosa: 1 9 (holotype), Taihorin, April, 1910, coll. H. Sauter (ZMB). This species seems to be closely related to P. macrodentatus (Van Lith, 1962) and therefore, although the enclosed area of the propodeum is not linear, I have placed it in the group of P. guadridentatus. Tsuneki (1966) described two males from Formosa, with somewhat paler legs, but otherwise very similar, as P. guadridentatus formosanus. As long as the shape of the clypeal margin of the corresponding Formosan female is unknown there remains some doubt in regard to the systematic status of formosanus and also the validity of the name parvidentatus. Group of Psenulus nasicornis Psenulus nasicornis spec. nov. Female. — Head black; mandibles except black teeth and scape of antennae yellow. Palpi, labrum and flagellum of antennae yellowish-red. Thorax black; dorsal part of pronotum and pronotal tubercles yellow. Scutellum and metanotum, except scutellar anterior groove and the depressed lateral parts, yellow. Axillae black. Fore and mid legs including coxae yellow, tarsi and hind legs completely yellowish-red. Veins of wings yellowish-red, subcosta brown. Gaster including petiole yellowish-red. Median part of clypeal margin much protruding and with two small teeth, close together (Fig. 12). Lower half of clypeal disk protruding medianly into a distinct tooth (Fig. 15). Puncturation of disk dense but not deep. Interantennal carina raised and much broadened between antennae (Fig. 13), dorsal surface hardly concave, ending 164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Fig. 9—15. Species of Oriental Psenulus. 9—11, head in frontal and dorsal view, pygidial area of female of P. parvidentatus sp. nov. (Formosa); 12—15, head, interantennal carina in dorsal view, pygidial area, face in lateral view of female of P. nasicornis sp. nov. (Celebes) J. P. van LITH: Oriental Psenulus 165 below antennae in an arched, distinct, transverse carina. Frons and vertex not distinctly punctate. Labrum with large emargination. Mandibles bidentate at apex and with a strong basal tooth. Scape of antennae concave on inner side, third antennal segment with deep groove on basal two thirds, also fourth segment somewhat depressed; third segment about two and a half times and fourth segment about one and a half times as long as broad at apex, segments 9—11 about as long as they are broad at apex, last segment about one and two-thirds times as long as broad at base. Anterior lateral corners of pronotum pointed. Scutum with very fine piliferous punctures and with many large but shallow punctures (interspaces between the latter once or twice diameter of punctures). Length of prescutal sutures just over one third of scutum. Scutellum and metanotum finely punctate. Enclosed area of propodeum de- pressed, posteriorly not sharply defined; also oblique longitudinal carinae unsharp. Enclosed area and longitudinal groove separated by a semicircular strong carina. Groove except widened upper part narrow, ending before apex in a few strong transverse carinae. Back of propodeum with wide-spread puncturation, lower half and sides of propodeum with transverse carinae, carination on the sides more irregular. Anterior part of sides of propodeum smooth. Mesopleura and metapleura almost smooth. Anterior oblique suture weakly foveolate, widened upper part with a few very weak transverse carinae. Apex of mid tibiae with a longitudinal row of four, sometimes five, reddish thorns and another single tooth at the apical margin. Base of hind tibiae on outer side with cluster of small red teeth, halfway tibiae one much stronger thorn, sometimes two thorns, and also apex with a few reddish thorns. Tibiae bare between the small basal teeth and the thorn half- way. First recurrent vein of fore wings about interstitial. Petiole cylindrical. Second gastral sternite without distinct semi-elliptic depression. Gaster with very fine piliferous punctures. Pygidial area narrow, lateral carinae almost parallel, only at base somewhat diverging (Fig. 14). Pubescence of face silvery and appressed, with a few long erect hairs. Hairs on rest of body yellowish. Back of propodeum fully pubescent, enclosed area bare. On lower part of epicnemial areas a large patch of yellowish appressed pubescence. Mesosternum with short dense pubescence and a number of long and erect hairs. Margin of fourth and fifth sternites with dense, short, white pubescence; disk of sixth sternite with dense, short, yellowish pubescence. Wings densely yellowish-pubescent. Length about 11,5 mm. Male unknown. SE Celebes: 3 9 (holotype and paratypes), Tangke Salokko Mountain, 1500 m, 1—15 Jan., 1932, coll. G. Heinrich (ZMB, one paratype in the author’s collection). This large wasp shows affinities with the group of P. guadridentatus, in some respects also with the group of P. pulcherrimus, but the combination of its characters does not allow its placing in any of these groups. The species is easily distinguished by the broad interantennal carina, which is not distinctly concave dorsally, and by the tooth on the clypeal disk. Group of Psenulus antennatus Psenulus bengalensis spec. nov. Male. — Head and thorax black, mandibles also dark, underside of first two segments of antennae brown, basal half of underside of segments 3—6 brown, palpi testaceous, 166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 labrum dark brown, pronotal tubercles dark brown, tegulae reddish-brown. Legs pale red, trochanters and underside of fore and mid femora brown. Gaster including petiole pale red. Veins of wings brown. Clypeal disk convex, more than in P. antennatus (Rohwer), anterior margin of clypeus with two short teeth (Fig. 16). Clypeus and face not distinctly punctate. Interantennal carina raised, broadened and concave between antennae (Fig. 17), ending below antennae in a transverse carina. Frons and vertex with minute punctures. Antennae moniliform, first segments of flagellum somewhat depressed. Antennal segments 3—12 about twice as long as broad at apex, segments 4—12 with distinct, narrow, oblique tyloidea, on fourth segment length of tyloidea about one-third of length of segment, on segments 5—8 about half that length, on ninth and 10th segment slightly shorter and on 12th segment very small, last segment narrower, more than twice as long as broad at base. Scutum with fine punctures, intermixed with a few large punctures, which are most distinct along the prescutal sutures. The latter sutures only distinct on anterior half, on posterior part replaced by an irregular row of large punctures. Anterior oblique suture coarsely foveolate, widened upper part with transverse carinae. Mesopleura with fine punctures. Hypo-epimeral area distinct. The scutellar suture is deformed in the holotype. Enclosed area of propodeum longer than in P. antennatus 8, with a distinct, slightly irregular median longitudinal carina. Upper two-thirds of back of propodeum (Fig. 18) ORO = | ea 16 17 OVO 2 A X, 19 20 I Fig. 16—20. Species of South-Asiatic Psenulus. 16—18, head, interantennal carina in dorsal view, propodeum of male of P. bengalensis sp. nov.; 19—20, head, interantennal carina in dorsal view of female of P. nietneri sp. nov. J. P. van LitH: Oriental Psenulus 167 smooth, longitudinal sulcus broad and deep, especially in the middle part of the propo- deum, above and below this deeper part a few transverse carinae. Between hind surface and sides of propodeum an edge of coarse, irregular carination, lower third of hind surface of propodeum also with coarse, irregular carination. Greater part of sides of propodeum smooth. Legs slender. Second submarginal cell normal. First recurrent vein of fore wings ending in second submarginal cell, second recurrent vein ending well in third sub- marginal cell. Petiole reaching about as far as hind femora. Anterior half of petiole almost quadrate, rest of petiole nearly cylindrical. Face with silvery pubescence, mostly appressed, rest of body with somewhat yellowish pubescence. Length about 7 mm. Female unknown. India: 1 & (holotype), Bengal, Senchal Range, Darjeeling, 27 July, 1923, coll. J. C. M. Gardner, No. 63 and No. 320 (USNM). P. bengalensis differs from P. penangensis (Rohwer) — judging from the original description of the female holotype — in having a smooth propodeum and a longer enclosed area of the propodeum. It is closely related to P. antennatus, but differing in the colour of the petiole, as well as in the hind surface of the propodeum being almost completely smooth. P. antennatus too has tyloidea on antennal segments 3—11, but these are indistinct. There are also close relationships with the group of P. quadridentatus, in which the petiole sometimes is red and the hind surface of the propodeum is smooth. However, in this group the enclosed area of the propodeum is much reduced and in the male of P. quadridentatus (Van Lith, 1962) only indistinct tyloidea can be found on segments 4—6. The pin also bears the labels “parasitic on” (printed) and “not yet id[enti}f[ie}d Hepialidae” (handwritten). Probably a hepialid caterpillar and the wasp were found together in a stem or twig, which is the usual nesting place of Psenulus. The prey of this species will undoubtedly prove to be a homopteron. Psenulus nietneri spec. nov. Female. — Head and thorax black; mandibles and labrum yellowish-brown; scape of antennae yellowish-red, upper side of flagellum dark brown, underside of flagellum reddish-brown. Pronotal tubercles, tegulae, legs including trochanters, and gaster including petiole yellowish-red. Coxae brownish. Veins of wings dark brown. Frontal carina between antennae raised, broadened and concave (Fig. 20), narrowed below antennae and ending in an arched tubercle which is not sharpened into a trans- verse carina. Anterior margin of clypeus with two distinct triangular teeth (Fig. 19). Frons densely and distinctly punctate, with fine oblique striation between anterior ocellus and oculi. Puncturation of vertex more wide-spread. Antennae somewhat clavate; third segment about twice as long and segments 9—11 about as long as broad at apex; last segment slightly more than one and a half times as long as broad at base. Prescutal sutures distinct on anterior two-thirds of scutum, vague on posterior part. Anterior third of scutum with fine transverse striation, in front of scutellum some irregular longitudinal striation. Scutellum and metanotum with fine and widespread puncturation. Enclosed area of propodeum normal, behind this a smooth and shining area; back of propodeum 168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 with fine sculpture and punctures. Median sulcus broad and deep on upper half, lower half narrower, almost reaching apex. Sides of propodeum smooth, between back and sides an irregularly carinate edge. Mesopleura densely but not deeply punctate. Anterior oblique suture coarsely foveolate, widened upper part with some transverse carinae. Second submarginal cell normal. First recurrent vein of fore wings interstitial, second recurrent vein ending well in third submarginal cell. Petiole cylindrical, reaching about as far as apex of hind femora. Second gastral sternite without distinct semi-elliptic depression. Pygidial area only indicated by a slight, narrow, longitudinal depression, no lateral carinae. Face with appressed silvery pubescence and also with long, erect, whitish hairs. Con- cave broadened part of interantennal carina laterally with a row of whitish hairs. Thorax with whitish pubescence, which is denser on mesosternum; epicnemial areas below with a round patch of yellowish pubescence. Gaster with yellowish-grey pubescence. Margin of fourth and fifth gastral sternites with dense and short white pubescence. Length about 7 mm. Male unknown. Ceylon: 1 9 (holotype), coll. Nietner (ZMB). This species is easily distinguished from P. antennatus by the red legs and petiole. Group of Psenulus erraticus Psenulus erraticus Smith 1861, Smith, Jl. Proc. Linn. Soc. Zool. 5: 85, ® (Psen erraticus; Celebes). 1962, Van Lith, Zool. Verh. Leiden 52: 48-49. 1970, Van Lith, Steenstrupia 1 (10): 95-96. New records: Vietnam: 2 9, Dai Lanh, N of Nha Trang, 30 Nov.—5 Dec., 1960, coll. C. M. Yoshimoto (BISH). These females have been compared with a female collected in the Philippine island Tawi Tawi and which probably belongs to the subspecies baszlanensis (Rohwer). They are all very similar. However, the back of the propodeum medianly, just behind the enclosed area, is not densely pubescent as in the Philippine specimen, but almost bare, leaving the coarse reticulate carination distinctly visible. The petiole is less heavy. They may belong to a new subspecies. Thus far I have seen females from Vietnam, Borneo, Celebes, Tawi Tawi and Basilan and males from Singapore and Mindanao. A comparative study, based on more extensive material, is needed. The characters used for distinguishing the subspecies are only slight and their constancy should be confirmed. Group of Psenulus crabroniformis Psenulus crabroniformis (Smith) 1858, Smith, Proc. Linn. Soc. Zool. 2: 107, “9” (4!) (Mellinus crabroniformis; Borneo). 1887, Handlirsch, Sitzb. kais. Akad. Wissensch. 96: 294. 1907, Cameron, Jl. Straits Br. R. As. Soc. 48: 23-25. 1962, Van Lith, Zool. Verh. Leiden 52: 51-53 (Psenulus crabroniformis). Two females from Thailand differ so little from the holotype ( g' ), from Borneo, that I am not able to distinguish these females as a new subspecies. The Thailand specimens J. P. van LITH: Oriental Psenulus 169 have darker femora, the greater part being dark brown instead of pale red, as in the male from Borneo, and the hind femora have small black streaks which are lacking in this male. The veins of the wings are dark brown; these are much paler in the holotype but they may have faded in this old specimen. Description. Female. — Head black with following exceptions: mandibles yellow with dark red tips, palpi light brown, labrum reddish, scape of antennae yellow, underside of flagellum reddish-brown. Thorax black, but pronotal tubercles yellow, tegulae pale red. Fore and mid legs yellow, including tips of coxae, but greater part of femora dark brown. Hind legs: tips of coxae reddish, trochanters yellow, rest of legs red except a small black stripe on fore side and back of femora. Gaster, including petiole, red, base of petiole paler. Veins of wings brown. Clypeus as in the female of P. crabroniformis sumatranus (Ritsema). Depressed margin with broad triangular emargination. Lower part of clypeal disk triangularly raised, indistinctly but densely punctate, upper third almost impunctate. Frontal carina prominent between antennae, dorsally slightly broadened, but not excavate. It ends in a transverse carina below antennae which is not much protruding and which does not reach the sides; laterally it is bent upwards to the antennal sclerites. Frons and vertex smooth and shining, punctures minute, a small depression on outer side of lateral ocelli. No tubercles on frons. Antennae rather slender, slightly clavate, segments 9—11 about as long as they are broad. Pronotum dorsally with sharp anterior carina, declining foreside of pronotum trans- versely striate. Scutum shining, with few punctures, prescutal sutures indistinct. Scutellum long and flat, scarcely punctate, with broad and foveolate scutellar suture. Metanotum posteriorly with very fine punctures. Enclosed area of propodeum triangular, depressed, with sharp, oblique, longitudinal carinae. Horizontal part of propodeum behind enclosed area smooth and shining. Posterior surface, except a very small area on both sides of median sulcus, and sides of propodeum coarsely reticulate, median upper part of propodeal declivity depressed. Upper half of median sulcus of propodeum normal, as broad as in P. crabroniformis sumatranus; lower half of sulcus widened towards apex, shining. Mesopleura and mesosternum sparsely and finely punctate, hypo-epimeral area convex, sparsely punctate. Tarsi of hind legs long, inner spur of hind tibiae about two- thirds of length of basitarsus. First recurrent vein of fore wings received by first, second recurrent vein by third, submarginal cell. Petiole long and slender, cylindrical, reaching beyond apex of hind femora. Gaster with minute punctures, shining, last two tergites very finely coriaceous. No distinct pygidial area. Appressed pubescence of face silvery. Pubescence of head, thorax and gaster yellowish- grey. Hind femora below with long hairs, upper half of back of femora bare. Lower part of epicnemial areas with round patch of dense pale pubescence. Pubescence of mesosternum long, densest along posterior part of median line. Length about 10 mm. Thailand: 2 9 (one allotype), Fang, 29 Nov., 1962, coll. A. Nagatomi (HUA). A female from the Philippine Is. is also very similar to specimens of the nominate form from Borneo. As in the Thailand females the wing-veins are dark brown. The female differs further from the Bornean male in the whitish pubescence of head and thorax. The gaster is lacking. Fore and mid femora light brown, also tips of coxae. Hind legs reddish, including tips of coxae, apex of hind tibiae brownish. Frons without distinct punctures. Antennal segments 9-11 slightly longer than they are broad. 170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 A study of the indices of the posterior interocellar distance and the ocellar-ocular distance in the various forms of P. crabroniformis might be interesting. In one of the Javanese females of P. crabroniformis sumatranus the index is 1: 134, in the female from Busuanga 1: 14. Philippine Is.: 1 9, Busuanga I., 4 km N of San Nicolas, 31 May, 1962, coll. H. Holt- mann, Malaise trap (BISH). Group of Psenulus tuberculifrons Psenulus tuberculifrons tuberculifrons (Rohwer) 1921, Rohwer, Phil. JI. Sc. 18 (3): 315-316, 2 (Diodontus tuberculifrons; Luzon). 1962, Van Lith, Zool. Verh. Leiden 52: 57-59 (Psenulus tuberculifrons tuberculifrons). New record: 1 ©, Luzon, Los Baños, 1917, coll. F. X. Williams (BISH). Psenulus cavifrons Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 60, 9 (Samar). New record: 1 9, Mindanao, Zamboanga del Norte, 9,6 km E of Sindangan, 20 July, 1958, coll. H. E. Milliron (BISH). Scutum with small yellow mark along tegulae (Fig. 21). A small yellow spot on anterior margin of scutellum. Anterior oblique suture narrow but foveolate, upper part widened and with indistinct transverse carinae. Second gastral sternite with long semi- elliptic depression, which is not distinctly defined. Pygidial area: Fig. 22. Mesosternum with long, backwards-directed, whitish hairs, brown medianly. Epi- cnemial areas below with round patch of dense pubescence. Margin of fourth and fifth gastral sternites with dense and short, whitish pubescence. Shields on outer side of lateral depressions of metanotum with a short black tooth, directed obliquely inwards (Fig. 21), which seems to be transition towards the long projections occurring in the group of P. nigrolineatus. I also noticed this structure in P. suluensis (Van Lith, 1970). It may be another proof of the close relationships between the group of P. twberculifrons (base of hind tibiae with three long spines and, in the Philippine species, with frontal tubercles) and the group of P. nigrolineatus (metanotum with very long, inwards-directed, projections, no frontal tubercles and only P. diversus having long tibial spines). Group of Psenulus nigrolineatus Psenulus diversus Van Lith 1923, Rohwer, Phil. Jl. Sc. 22 (6): 597, ® (Diodontus ajax partim; Singapore). 1962, Van Lith, Zool. Verh. Leiden 52: 70-71 (Psenulus diversus). A female from Vietnam is somewhat darker than the holotype from Singapore. Median dark line on scutum broader on anterior two-thirds and connected with posterior margin by a narrow line (Fig. 26). Posterior surface of propodeum yellow, divided into four longitudinal parts by two black lines. Hind margin of scutellum more J. P. van LITH: Oriental Psenulus 171 Fig. 21—26. Species of Oriental Psenulus. 21—22, thorax and pygidial area of female of P. cavi- frons Van Lith (Mindanao); 23, scutum of male of P. luteopictus luteopictus (Rohwer) (Luzon); 24—25, genitalia in dorsal and ventral view of male of P. sandakanensis (Rohwer) (Borneo); 26, thorax of female of P. diversus Van Lith (Vietnam) extensively black. Upper part only of anterior plate of mesepisternum and anterior two- thirds of hypo-epimeral area yellow. Scape of antennae and basal two-thirds of flagellum yellow. Fore femora with a dark brown mark on posterior side, mid femora with yellow foreside and reddish hind surface with yellow knee, hind femora reddish brown, basal half of hind surface yellow. Petiole not darkened at apex, in the holotype the apex is slightly darkened. Third segment of antennae about three times, fourth and fifth segments slightly more than twice as long as broad at apex, following segments gradually shortening, segments 172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 9—11 about one and a half times as long as broad at apex, last segment nearly twice as long as broad at base. Anterior oblique suture foveolate, widened upper part smooth. First recurrent vein of fore wings ending in first submarginal cell, second recurrent vein ending in third sub- marginal cell. Second gastral sternite without distinct semi-elliptic depression. Margin of fourth and fifth sternites with dense, short and pale pubescence. Vietnam: 1 9, Dai Lanh, N of Nha Trang, 30 Nov.—5 Dec., 1960, coll. C. M. Yoshimoto (BISH). Notwithstanding the long spines at the base of the hind tibiae, which are absent in the other forms of the group of P. nigrolineatus, I prefer to maintain P. diversus provision- ally in this group. The form from Vietnam may belong to a different subspecies. Group of Psenulus luteo pictus Psenulus luteopictus luteopictus (Rohwer) 1921, Rohwer, Phil. Jl. Sc. 18 (3): 319-320, 2 (Diodontus luteopictus; Luzon). 1962, Van Lith, Zool. Verh. Leiden 52: 71-72, 2 (Luzon, Negros). Male. — Head black; mandibles yellow except reddish and black tips, palpi yellowish- brown, scape of antennae and first segment yellow, following segments yellow below, dorsally brown. Ventral and lateral sides of thorax black; yellow are: dorsal side of pronotum, tuber- cles, scutum except narrow antero-lateral black margin, a large median, oval, black spot, connected with fore margin by a broad line, two triangular black marks on outer sides of prescutal sutures (Fig. 23) and a narrow black hind margin of scutum. Scutellum including axillae and metanotum yellow, depressed lateral parts black. Propodeum yellow; greater part of enclosed area and two adjacent lateral spots, longitudinal stripe over upper half of sulcus on back of propodeum, and apical margin black. Upper two-thirds of anterior plate of mesepisternum, hypo-epimeral area and small adjacent spot below, and upper part of metapleura also yellow. Subalar area of fore wings black. Fore and mid legs including trochanters and lower third of coxae yellow; hind legs brownish but coxae more yellow and fore side of femora darker brown. Veins of wings brown. Gaster including petiole reddish, apical spine reddish with dark tip. Clypeal margin bidentate, on lower third of clypeus, between the teeth, a semi-elliptic depression. Clypeus and rest of face shining, with fine puncturation. Frontal carina raised into a sharp plate between antennae, not broadened, margin somewhat transparent. Frontal carina ending in a semicircular transverse carina. Frons and vertex shining, with dense piliferous puncturation. Interocellar region raised, bare and shining with only a few irregular punctures, therefore conspicuous as compared with the densely pubescent frons and vertex. Antennae long and slender, slightly clavate, third segment at least three times, fourth segment nearly three times as long as broad at apex, 10th and 11th segments about one and one-third and 12th segment one and a half times as long as broad at apex; last segment about twice as long as broad at base. Pronotal corners sharp. Scutum with strong puncturation, coarse centrally, interspaces there partly smaller than diameter of punctures. Scutum along basal part of parapsidal sutures somewhat depressed. Length of prescutal sutures about one fourth of scutum. J. P. van LitH: Oriental Psenulus 173 Enclosed area of propodeum not sharply defined behind, oblique carinae exceeding black part of enclosed area. Back and sides of propodeum with coarse reticulate carination. Median longitudinal sulcus narrow, reaching about halfway back of propodeum. Anterior oblique suture simple, widened upper part smooth. Mesopleura sparsely punctate. Legs long and slender. First recurrent vein of fore wings ending in first sub- marginal cell, distance from second submarginal cell about one third of base of second cell; second recurrent vein ending in third submarginal cell, distance from second sub- marginal cell about one fourth of base of second cell. Gaster slender, petiole long and surpassing end of hind femora, second sternite without semi-elliptic depression. Last tergites somewhat depressed laterally. Face including frons and also vertex densely pale-golden pubescent, appressed below transverse carina. Tempora with short silvery pubescence and few long hairs. Hairs on back of head, and on thorax, long and yellowish. Pubescence on gaster short, but petiole with a few long lateral hairs. Length nearly 8 mm. Philippines: 1 & (allotype), Luzon, Mt. Montalban, Rizal, Wa-wa Dam, 150—200 m, 8 March, 1965, coll. H. M. Torrevillas (BISH). The male of P. /wteopictus, now described for the first time, very much resembles the females from Luzon, even in the golden pubescence of the face. The strong puncturation of the scutum seems to be somewhat more dense. Group of Psenulus sandakanensis Psenulus sandakanensis (Rohwer) 1923, Rohwer, Phil. Jl. Sc. 22 (6): 597-598, & (Diodontus sandakanensis). 1962, Van Lith, Zool. Verh. Leiden 52: 73-76, & and ® (Psenulus sandakanensis). 1967, Van Lith, Ent. Ber. 27: 20, 4. New records: N Borneo (SE), Forest Camp, 19 km N of Kalabakan, 60 m, 2 9, 17 Oct., 1962, coll. K. J. Kuncheria; 1 9 and 1 &, 28 Oct, 1962 and 1 G', 30 Oct. 1962 (Primary Forest), coll. Y. Hirashima (BISH). The females have no transverse carina below the antennae. Third antennal segment slightly more than twice, fourth and fifth segments about twice, and sixth and seventh segments about one and a half times as long as broad at apex, segments 8—11 some- what longer than broad at apex, and last segment about twice as long as broad at base. The anterior plate of the mesepisternum is black. Base of hind tibiae flattened and smooth, upper part of this flattened area margined by a number of tiny thorns. Margin of fourth and fifth gastral sternites with dense and short, whitish pubescence. In the males antennal segments 3—6 slightly more than twice, and segments 7—12 about twice as long as broad at apex, these latter segments slightly rounded below. Last segment nearly three times as long as broad at base, laterally flattened towards apex. No distinct tyloidea. Genitalia: Fig. 24 and 25. One of the above mentioned males has three black marks on the scutum instead of one and resembles the male from Bettotan (near Sandakan) recorded in 1962. Its mesopleura are partly black along the metapleura. The anterior plate of the mesepisternum is completely dark, while in the males from Sandakan the upper part is yellow. Sometimes the basal half of the first gastral tergite is darkened in the males. 174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 In both sexes the subalar area of the fore wings is much raised, its posterior half is yellow. The anterior oblique suture is narrow, but distinctly foveolate, the widened upper part is smooth. The first recurrent vein of the fore wings ends distinctly in the first sub- marginal cell, the veins are brown. Psenulus pagdeni Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 81-83, 2 (Malaya). New records: N Borneo (SE), Forest Camp, 19 km N of Kalabakan, 60 m, 2 Q, 25 Oct., 1962, coll. K. J. Kuncheria and 1 2, 1 Nov., 1962, Malaise trap, coll. Y. Hi- rashima (BISH). The apex of the petiole of the Bornean females is darker than in the Malayan speci- mens, the apical third part of petiole being black-brown, ventrally and dorsally, and also the basal two-thirds of ventral plate. In the holotype only the base of the ventral plate is darkened, in the paratypes the petiole is dorsally slightly or not darkened. The veins of the wings are brown. The third antennal segment is about two and a half times as long as broad at apex. Segments 9—11 are about as long as broad at apex, the last segment is about one and a half times as long as broad at base. The longitudinal groove on the back of the propodeum is narrow. The oblique anterior suture of the mesopleura is distinctly foveolate, the widened upper part is smooth. Group of Psenulus esuchus Psenulus esuchus (Rohwer) 1923, Rohwer, Phil. Jl. Sc. 22 (6): 600-601, & (Diodontus esuchus). 1962, Van Lith, Zool. Verh. Leiden 52: 79-80, 4 (Psenulus esuchus). 1967, Van Lith, Ent. Ber. 27: 20, 3. 1970, Van Lith, Steenstrupia 1: 100, 4 (Balabac). Female. — Head black; mandibles yellow with dark tips, palpi yellow, labrum reddish, anterior margin of clypeus somewhat brownish. Scape of antennae yellow, pedicel yellow below, reddish above. Pronotal tubercles yellow. Scutum (Fig. 27) yellow with three broad, black lines; lateral lines not reaching anterior margin of scutum, posteriorly often reaching hind margin; median mark as long as anterior three fourths of scutum, with narrow basal part. Tegulae yellow. Scutellum yellow, rarely with a vague and narrow, longitudinal, brown line. Axillae and metanotum yellow. Base of propodeum with broad black band, median longitudinal groove black, apex of propodeum with two black spots. Epicnemial areas except a yellow margin — in the closely related corporaali from Suma- tra only the outer keel is yellow — and interepicnemial area black. Anterior plate of mesepisternum yellow, hypo-epimeral area yellow except small, black, posterior tip. Anterior part of mesopleura yellow, connected with lower part of posterior margin by a yellow band. Metapleura black. Mesosternum black, roundly dilated on to lower part of sides of thorax. Fore and mid legs including coxae yellow, tarsi darker. Hind coxae yellow, rest of hind legs yellowish-red but femora dorsally, apical part of tibiae, and tarsi dark brown. Veins of wings brown. Petiole pale yellow, apex laterally dark brown, rest of gaster pale red. J. P. van LITH: Oriental Psenulus 175 Median part of anterior margin of clypeus straight, with two small teeth, broad depression behind margin, no distinct large punctures. Disk of clypeus strongly convex. Raised part of interantennal carina narrow, ending below antennae in a transverse carina, which is distinct but low. Frons and vertex with fine punctures. Antennae slightly clavate, third segment nearly two and a half times and segments 9—11 nearly one and a half times as long as broad at apex, last segment about twice as long as broad at base. Pronotum with sharp antero-dorsal carina. Scutum smooth, finely punctate. Prescutal sutures distinct, as long as anterior quarter of scutum. Enclosed area of propodeum triangular, median two carinae diverging; median longitudinal groove on posterior side of propodeum narrow, before apex ending in a transverse carina. Propodeum smooth behind enclosed area and behind metapleura, posterior surface with fine oblique rugae and fine reticulate carination, especially on the sides. Anterior oblique suture indistinctly foveolate, widened upper part smooth. Hypo-epimeral area well-defined. Mid tibiae with low longitudinal depression on apical fifth, defined behind by a row of four small red spines, anteriorly by an oblique raised area and an apical spine. Hind tibiae without distinct depression at base, outer side of base with a number of minute red tubercles. First recurrent vein of fore wings ending in first submarginal cell, rarely interstitial (sometimes in one wing only). Second gastral sternite with indistinct semi-circular depression. Pygidial area narrow, carinae parallel on apical half. Pubescence of head silvery, appressed on face, whitish on thorax, yellowish on gaster. Hind margin of fourth and fifth gastral sternites with dense and short pale pubescence, also over disk of sixth gastral sternite. Length 8—8,5 mm. New records from N Borneo: 12 9 and 1 g', Forest Camp, 19 km N of Kalabakan, 60 m, 15—30 Oct. and 4—19 Nov., 1962, coll. Y. Hirashima and K. J. Kuncheria (BISH); 1 9, Silabukan Forest Reserve, 1 Dec., 1968, coll. C. G. Roche (coll. Roche). In my opinion P. esuchus and P. corporaali (Van Lith, 1962) are very closely related, and probably the latter is a subspecies of the former. The main differences — in the females — are that the lateral black lines on the scutum are shorter in P. esuchus, but its hind legs and the apical part of the petiole are darker. The male of P. corporaali is still unknown. Because of the fine transverse carina below the antennae I have now placed P. esuchus, formerly incorporated in the group of P. interstitialis, into a separate group, together with a few closely allied species. Psenulus corporaali Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 76-77, 2 (Sumatra). Complementary description. Scutum: Fig. 28. Epicnemial areas laterally with narrow yellow margin. Subalar area of fore wings dark brown with small yellow tip. Yellow spot below hind wings. Tibial spines yellowish-brown. Third and fourth segments of antennae nearly two and a half times as long as broad at apex, following segments gradually shorter, 10th and 11th segments about one and a half times as long as broad at apex, 12th segment almost two and a half times as long as broad at base. Widened upper part of anterior oblique suture with indistinct transverse carinae. 176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Fig. 27—30. Species of Oriental Psenulus. 27, scutum of female of P. esuchus (Rohwer) (Borneo); 28, scutum of female of P. corporaali Van Lith (Sumatra); 29—30, thorax and pygidial area of female of P. varius Van Lith (Malaya) Tempora with mostly appressed silvery pubescence. Hairs on thorax more greyish than yellowish, long on propodeum, dense on mesosternum; epicnemial areas below with a round patch of dense, yellowish-grey pubescence. Pubescence on gaster yellowish, on margin of fourth and fifth sternites paler. Petiole with usual long, erect, hairs on sides. As regards relationships, see remarks on P. esuchus. Psenulus varius Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 69-70, & (Malaya). Two Psenulus females from Penang are supposed to belong to P. varius, but a male from this island should confirm the identification. They are very similar to the male, differing only in a few minor details. Also third antennal segment yellow below. Marking of scutum almost the same (Fig. 29). Scutellum darker, base with black triangle pointing backwards, in one female somewhat reduced. Apparently this triangle is so much reduced in the males that only a small black spot on the posterior half is left. Only dorsal part of propodeum black, back except median groove yellow. Subalar area of fore wings black, below hind wings a small yellow spot. J. P. van LitH: Oriental Psenulus 177 A large round mark on anterior half of hypo-epimeral area yellow. A large yellow mark on mesopleura behind anterior plate. Fore and mid legs yellow. Fore side of hind femora brown. Apex of petiole dorsally not darkened. Raised interantennal carina ending below antennae in a fine but distinct transverse carina. Antennae slender, third segment nearly three times, following segments about twice as long as broad at apex, gradually shortening, segments 10—11 about one and a half times as long as broad at apex, last segment about two and a half times as long as broad at base. Back of propodeum with fine carination, on lower two thirds irregularly reticulate, on upper part almost obliquely striate. Apex of mid tibiae with a short oblique ridge behind which a small depression and three or four short red thorns. Second gastral sternite with indistinct triangular depression. Pygidial area narrow, lateral carinae almost parallel (Fig. 30). Face with appressed silvery pubescence and also with many long silvery hairs. Epi- cnemial areas below with round patch of short yellowish-golden pubescence. Mesosternum with short yellowish-golden pubescence along median carina and also with dense, long and erect, whitish pubescence. Petiole with long and erect lateral hairs. Fourth and fifth gastral sternites, especially on apical margin, with dense and pale pubescence, sixth sternite with dense and short golden pubescence. Length about 8,5—9 mm. Malaya: 2 9 (allotype, 21 Jan., 1964), Penang, Tanjong Bungah, Sungei Kechil, 21 and 22 Jan., 1964, one with prey: a derbiid nymph (det. H. T. Pagden), coll. H. T. Pagden (coll. Pagden). The male was provisionally placed in the group of P. nigrolineatus, but judging from the female characteristics and the lack of the metanotal projections in both sexes, this species seems to be more closely related to P. eszchus. Psenulus pallidicollis spec. nov. Female. — Head black; mandibles pale yellow, with reddish tips, palpi pale yellowish- brown, scape of antennae yellowish-white, partly brown above, flagellum dorsally black, underside brown. Thorax black except following yellowish-white parts: dorsal side of pronotum, pronotal tubercles, two parallel longitudinal marks on posterior half of scutum (Fig. 31), axillae, lateral wings of scutellum, and metanotum except the depressed lateral parts and a small brown mark on its hind margin. Tegulae reddish. Fore and mid legs including trochanters yellowish-white, underside of femora brownish and tarsi reddish. Hind legs including coxae reddish, tarsi brown. Veins of wings brown. Posterior surface of propodeum with four elongate, yellowish-white, marks. Gaster, including petiole and apical spine, reddish. Clypeus convex, anterior margin depressed, indistinctly bidentate, distance between teeth about one fourth of total distance between eyes. Clypeal disk shining, with irregular puncturation. Raised interantennal carina narrow and sharp, ending below antennae in a fine transverse V-shaped carina. Frons and vertex smooth and shining, with fine piliferous punctures. Occipital carina normal. Apex of mandibles bidentate. Antennae long, third segment about three times as long as broad at apex, segments 4 and 5 about two and a half times and following segments about twice as long as broad at apex, last segment more than twice as long as broad at base. 178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 32 33 Fig. 31—33. Thorax, second and third submarginal cells of right fore wing, pygidial area of female of P. pallidicollis sp. nov. (Borneo) Scutum finely and sparsely punctate, prescutal sutures distinct only on anterior third of scutum. Parapsidal sutures sharp. Enclosed area of propodeum concave, with oblique carinae, median depression heart-shaped. Longitudinal median groove on back of propodeum reaching about half-way, lower three-fourths of back of propodeum and posterior part of sides of propodeum with reticulate carination. Anterior oblique suture rather broad, especially lower part with strong and relatively long transverse carinae, upper part widened with a few weak transverse carinae. Apical third of mid femora posteriorly with four short reddish thorns, anteriorly with a reddish, short and oblique, ridge and one short thorn at apex. Base of hind tibiae with a cluster of tiny reddish thorns. First recurrent vein of fore wings just ending in second submarginal cell, second recurrent vein ending in third submarginal cell (Fig. 32). Petiole cylindrical, long, reaching about as far as apex of hind femora, apex dorsally with a small triangular depression. Gaster very finely punctate. Pygidial area narrow, fine lateral carinae for greater part parallel (Fig. 33). Face and tempora with silvery, appressed pubescence and also many long, raised, J. P. van LITH: Oriental Psenulus 179 whitish hairs. Rest of head and thorax with long, whitish pubescence, somewhat yel- lowish on scutum, and dense on mesosternum. Epicnemial areas below with large, yellowish pubescent, circular area. Pubescence of gaster yellowish, posterior margin of fourth and fifth gastral sternites with dense and short, pale pubescence and sixth sternite with dense, backwardly directed and longer, golden pubescence. Length about 11 mm. Male unknown. Borneo: 1 Q (holotype), Sarawak, Mt. Dulit, 4000 ft, Moss Forest, 29 Oct., 1932, Oxford Univ. Exp., B. M. Hobby and A. W. Moore (BM, 1933— 254). Apparently this wasp is closely related to P. esuchus, having a fine transverse carina below the antennae. It differs from the other species belonging to the group of P. esuchus in the darker scutum and the yellowish-white colour of the markings of thorax and legs. Psenulus neptunus spec. nov. Female. — Head black, mandibles yellow with dark tips, palpi yellowish-brown, anterior margin of clypeus dark reddish-brown, scape and first antennal segment yellow, flagellum pale reddish. Thorax yellow, with following black parts: three irregular longitudinal black marks on scutum, about as long as posterior three quarters and con- nected along posterior margin (Fig. 36), scutellar suture, enclosed area of propodeum and a small adjacent black mark on the sides, median longitudinal sulcus of propodeum and lower margin of back of propodeum, a black band on fore side of pronotum, in- terepicnemial area, mesosternum partly extending on lower part of mesopleura, a mark along posterior margin of mesopleura, and metapleura. Fore and mid legs including coxae yellow, tarsi somewhat reddish; hind legs reddish with yellow coxae. Gaster in- cluding petiole pale reddish. Veins of fore wings, except subcostal vein, yellowish-brown. Disk of clypeus convex, anterior margin depressed, this depression about as broad as one fourth of length of clypeus, margin with two distinct teeth, distance between these teeth about one-fifth of total distance there between the eyes. Frontal carina raised, between antennae not broadened, ending in an arched transverse carina below antennae, about half as long as distance between the eyes (Fig. 34). Apex of mandibles bidentate. Antennae not clavate, third segment about two and one-third times and segments 9—11 about one and a half times as long as broad at apex, last segment twice as long as broad at base. Frons with very fine punctures from which fine appressed hairs arise, and with two converging lines each consisting of three strong punctures bearing a long hair, in front of anterior ocellus. Vertex with a few distinct punctures. Scutum with fine puncturation, intermixed with sparse, much larger, punctures. Scutellum and metanotum with fine punctures. Lateral depressions of scutellum and metanotum normal. Enclosed area of propodeum triangular, depressed, with oblique carinae. Back of propodeum with distinct oblique striation, upper part behind enclosed area almost smooth, sides with irregular reticulate carination. Anterior oblique suture weakly foveolate, widened upper part smooth. Legs normal, mid tibiae at apex with almost triangular flattened area bordered anteriorly by an oblique ridge, posteriorly by a few reddish spines and with two short spines at apical margin. Hind tibiae with a few reddish spines on apical margin. Second submarginal cell normal, first recurrent vein of fore wings almost interstitial, second recurrent vein ending in third submarginal cell (Fig. 35). Petiole cylindrical. 180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 35, Fig. 34-36. Head, second and third submarginal cells of left fore wing, scutum of female of P. neptunus sp. nov. (Celebes) Second gastral sternite without well-defined semi-elliptic depression. Pygidial area long and narrow, lateral carinae distinct. Face and frons with pale golden, dense and short, not appressed pubescence, leaving sculpture visible, and with many long and erect hairs. Head and thorax with long greyish-golden hairs, gaster with shorter greyish-golden hairs. Epicnemial areas with a round patch of dense and short greyish-golden pubescence. Apical margin of fourth and fifth gastral sternites with dense, short, pale pubescence. Sixth sternite with dense, short, golden pubescence. Wings with yellowish-grey pubescence. Length about 10 mm. Male unknown. Celebes: 1 9 (holotype), Ile Ile, 500—800 m, early Dec., 1930, coll. G. Heinrich (ZMB). This species has been named neptunus because of the tridentate marking of the scutum. P. neptunus and P. segrex, both from Celebes and both with pale wing-veins, are J. P. van LITH: Oriental Psenulus 181 similar in many respects. They belong to different species groups, however, as the transverse carina below the antennae is lacking in the female of P. segrex. Moreover, the pubescence of the face and the black marking of the scutum also differ. Group of Psenulus pseudajax Psenulus pseudajax holtmanni subsp. nov. Two males from the Philippines differ slightly from the male of the nominate form from Central Java. The yellow marks of the scutum, along the tegulae, are longer but they do not reach the anterior margin of the scutum (Fig. 37). Scutellum black with yellow hind margin, in the male from Culion the yellow marking is more extended (Fig. 37). The four yellow marks on propodeum confluent on lower part (Fig. 38), whilst in the nominate form they are fully separated. Fore and mid legs completely yellow, only in the male from Busuanga base of fore femora with a small brown spot. Gastral sternites darkened. Genitalia: Fig. 39 and 40. Pubescence and structure of head as in the male from Java. Antennae long, third segment about three and a half times as long as broad at apex, following segments gradually shorter, segments 10—12 nearly twice as long as broad at apex, last segment more than twice as long as broad at base. No tyloidea. SR Fig. 37—40. Thorax, propodeum, genitalia in dorsal and ventral view of male of P. pseudajax holt- manni subsp. nov. (Culion) 182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Anterior oblique suture indistinctly foveolate, widened upper part with some weak transverse carinae. Anterior margin of scutum with a few large punctures. As in the male from Java the first recurrent vein of the fore wings is interstitial, the metanotum has the same peculiar, long and narrow projections and the prescutal sutures are lacking. This latter characteristic is unusual in Psenulus. P. pseudajax certainly does not belong to the group of interstitialis, where it was provisionally placed in 1962, and has now been brought into a separate group. Philippine Is.: 1 _& (holotype), Culion I., 6 km W of Culion, 9 June, 1962, Malaise trap; 1 G', Busuanga I., 4 km N of San Nicolas, 23—24 May, 1962, Brushes, both coll. H. Holtmann (BISH). The male from Busuanga is somewhat darker, the scutellum being black with only a narrow yellow hind margin and the yellow marks on the propodeum are scarcely con- fluent. Group of Psenulus interstitialis Psenulus interstitialis interstitialis Cameron 1906, Cameron, Tijdschr. v. Ent. 49: 222-223, ® (Psenulus ? interstitialis, New Guinea). 1962, Van Lith, Zool. Verh. Leiden 52: 84-86, 9 (Psenulus interstitialis interstitialis). The females of P. interstitialis, recorded below, certainly belong to the nominate sub- species. They have all a completely yellow scutum, the metapleura are yellow, the extent of the black parts of the lateral depressions of the scutellum is variable, the enclosed area of the propodeum and the longitudinal sulcus are yellow or more or less black, and the median groove of the mesosternum is yellow, rarely with a complete black line. Anterior margin of clypeus reddish (Fig. 41, holotype). Veins of wings yellowish, but subcostal vein brownish. Antennae yellowish-red, dorsally brownish; third antennal segment about two and a half times, fourth and fifth segments about twice, sixth segment about one and three-quarters and segments 7—11 about one and a half times as long as broad at apex; last segment slightly more than twice as long as broad at base. Back of propodeum with fine oblique striation on upper part. Pygidial area: Fig. 42. Male. — Differs little from female, but median groove of mesosternum usually black. Dorsal side of antennae pale or darker brown. Distance between teeth on anterior margin of clypeus nearly one fifth or one sixth of the distance separating the eyes at that point. Interantennal carina ending below antennae in a distinct transverse carina, which is bilobed in frontal view. Antennae longer than in female, but scape shorter. Third antennal segment more than two and a half times, following segments about two and a half times as long as broad at apex; 13th segment about three times as long as broad at base; segments 8—12 somewhat rounded below, no distinct tyloidea. Posterior surface of propodeum and posterior part of sides of propodeum with coarse reticulate carination. Anterior oblique suture narrow, weakly foveolate, widened upper part smooth. Pubescence of frons and vertex pale golden, below antennae silvery or very pale golden and appressed, also with many long erect hairs. Genitalia: Fig. 43. Length about 10 mm. J. P. VAN LITH: Oriental Psenulus 183 45 A, S — 49 48 su Fig. 41—51. Subspecies of Psenulus interstitialis Cameron. 41—42, clypeus and pygidial area of female (holotype) of nominate form from New Guinea; 43, genitalia of male of nominate form (E New Guinea); 44—46, scutum, pygidial area, clypeus of female of P. interstitialis lutescens (Turner) (NE Australia); 47—51, genital apparatus of male of P. interstitialis salomonensis subsp. nov. (Solomon Is.), 47, dorsal view (Guadalcanal), 48—49, dorsal and lateral view (Russell Is.), 50—51, dorsal and ventral view (Bougainville) New records: NW New Guinea: 1 ¢ (allotype), Humboldt Bay, Hollandia, April, 1936, coll. L. E. Cheesman (BM, 1936—271); 1 9, Navire, S. Geelvink Bay, 1—20 m, 2—9 July, 1962, J. L. Gressitt and J. Sedlacek, Malaise trap (BISH); 1 9, Sorong, Kampong Malano, 28 Aug.—6 Sept., 1948, coll. M. A. Lieftinck (ML). Central N New Guinea: 5 9, Idenburg River, Bernhard Camp, 50 m, July—Nov., 1938, coll. J. Olthof, Neth. Ind.-American New Guinea Exp. (ML and USNM). 184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Territory of New Guinea: 1 9, Finschhafen, 6 Oct., 1944, coll. E. S. Ross (CAS); 1 ® and 2 G', Huon Peninsula, Finschhafen, 80 m, 16 April, 1963, coll. J. Sedlacek (BISH); 1 9, Nadzab, 25 June, 1944, coll. U. N. Lanham (Coll. Krombein); 1 9, Nadzab, Markham River val., along east fork of Ngafir Creek, 13 Aug., 1944, coll. K. V. Krombein (Coll. Krombein). Papua: 1 @, Kokoda, 1200 ft, April, 1933, coll. L. E. Cheesman (BM); 1 9, Cape Rodney, 2 Nov., 1960, coll. J. L. Gressitt, Malaise trap; 1 9, Daru I., 2 m, 24—25 Oct. 1960, coll. J. L. Gressitt, Malaise trap; 1 9, Owen Stanley Range, Goilala, Loloipa, 16—30 Jan., 1958, coll. W. W. Brandt; 1 4, W District, Oriomo Govt. Station, 26—28 Oct., 1960, J. L. Gressitt, Malaise trap (BISH). Some females and males from New Guinea, even those collected at the same locality as the normal form, and also two females from Japen Island, are darker than the holo- type. Scutum with black mark in the middle, metapleura and median groove of mesoster- num more or less black. Enclosed area of propodeum and median sulcus on back of propodeum almost completely black. The antennae of the females are of normal length. Provisionally, I consider these darker specimens as a variety of the nominate form. The black mark on the scutum is similar to that of the NE Australian subspecies /utescens Turner, but in the latter form the metapleura are completely yellow, at least in the four females which I have studied. Moreover, the distance between the teeth on the clypeal margin is larger, about one-fifth of the distance separating the eyes at that point, while in /utescens it is about one-seventh. NW New Guinea: 1 4, Nabire, S Geelvink Bay, 16 Sept., 1962, in jungle, coll. H. Holtmann, Malaise trap (BISH). Japen Island: 2 ©, Mt. Baduri, 1000 ft, Aug., 1938, coll. L. E. Cheesman (BM, 1938—593). Territory of New Guinea: 1 G', Finschhafen, 15 April, 1963, coll. J. & M. Sedlacek, 1 9 and 1 &, Wau, 1250 m, 4 May, 1965, coll. J. Sedlacek; 1 &, Wau, 1200 m, 10 Sept., 1964, coll. J. & M. Sedlacek; 1 g, Wau, 1190 m, 11 Sept., 1964, coll. M. Sed- lacek, Malaise trap; 1 g', Karimui, 1080 m, 8—13 July, 1963, coll. J. Sedlacek, light trap (BISH). SE New Guinea: 1 g', Popondetta, 60 m, 30—31 Aug., 1963, coll. J. Sedlacek, Malaise trap (BISH); 1 G', Kapakapa, May—June, 1891, coll. L. Loria (MCG). Psenulus interstitialis lutescens (Turner) 1907, Turner, Ann. Mag. Nat. Hist. 19 (7): 273, ® (Psen lutescens; Australia, Queensland). 1908, Turner, Proc. Zool. Soc. London 30: 463 (Psenulus interstitialis). 1916, Turner, Ann. Mag. Nat. Hist. 17 (8): 128 (Neofoxia interstitialis). A year after its publication P. /wtescens was considered by its author as a synonym of P. interstitialis Cameron. I have seen the type (No. 21.838) from Mackay and also three females from Cairns, all from the collection of Turner in the British Museum (Natural History) and there is no doubt that they are closely related to P. interstitialis from New Guinea. Although the subspecies of P. interstitialis are all somewhat variable, I provisionally consider /wtescens as a distinct subspecies. The distance between the two teeth on the anterior margin of the clypeus is smaller, viz about one-seventh of the total distance there between the eyes (Fig. 46), as against about one-fifth in the nominate subspecies. J. P. van LITH: Oriental Psenulus 185 Scutum with large square mark in the middle (Fig. 44), scutum of nominate subspecies being usually completely yellow. Lateral depressions of scutellum for the greater part black. Metapleura yellow, as in the nominate subspecies. No broad black line on mesoster- num. Enclosed area of propodeum not fully black, in the holotype even completely yellow. Veins of wings yellowish. Pygidial area: Fig. 45. Length about 9,5 mm. Male unknown. NE Australia: 1 9 (holotype), Mackay, Dec., 1900 (12.00); 2 9, Cairns, Kuranda, Nov., 1901 (11.01) and Febr., 1902 (2.02), Turner Coll. 1909—49; 1 9, Kuranda, 1100 ft, 30 May, 1913, coll. R. E. Turner 1913—438 (BM). More material should be studied to find out whether the smaller distance between the clypeal teeth and the black marking of the scutum are constant characteristics. A form from New Guinea with similar marking on the scutum is perhaps a variety of the nominate subspecies. It differs from /ztescens not only by the larger distance between the clypeal teeth, but also by its darker metapleura. Psenulus interstitialis salomonensis subsp. nov. The form of P. interstitialis from the Solomon Islands differs from the New Guinean forms in the yellowish-red, instead of yellow, colour of the thorax. In the female also the fore and mid legs are darker. Only the mandibles and the scape of the antennae are paler yellow. Flagellum of antennae yellowish-brown, in the male somewhat darkened apically. Scutum without black mark in the middle. Metapleura pale yellowish-red in female, metapleura of male either pale yellowish-red or more or less black. Extent of black colour in enclosed area of propodeum somewhat variable. Depressed lateral parts of scutellum black, of metanotum yellow. Median groove of mesosternum more or less black. Hind margin of scutum of male black. Veins of wings yellowish-brown. In both sexes frons and vertex flatter than in the New Guinean and NE Australian forms. Distance between clypeal teeth about one-fifth of total distance there between the eyes. Anterior oblique suture almost linear, widened upper part smooth. Face with silvery appressed pubescence and also a number of long, erect, hairs. Pubescence of rest of body yellowish-brown. Female. — Upper part of posterior surface of propodeum almost smooth, indistinctly transversely striate. Antennae similar to those of the nominate subspecies. Third segment about two and a half times as long as broadest part at apex, segments 4—6 about twice as long, segments 9—11 about one and a half times as long as broad at apex, last segment about twice as long as broad at base. Posterior margin of fourth and fifth gastral sternites with short and dense whitish pubescence. Length about 8—9 mm. Male. — Posterior surface of propodeum with coarse and irregular carination. Anten- nae long, as in the nominate form, but last segment slightly shorter, less than three times as long as broad at base. Segments 8—11 slightly rounded below, no distinct tyloidea. Genitalia: Fig. 47—51. Length about 7,5—8 mm. Solomon Is.: 1 9 (holotype) and 1 & (allotype). Guadalcanal I., Tenaru R., Jan., 186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 1945, coll. G. E. Bohart (CAS); 1 9, Tambalia, 30 km W of Honiara, 22 May, 1964, coll. R. Straatman (BISH). Malaita I.: 1 G', Dala, 50 m, 6—13 June, 1964, coll. J. and M. Sedlacek, Malaise trap (BISH). Russell Id.: 3 © and 1 Z, Pavuvu I., Pepesala, O—100 m, 18—20 July, 1964, coll. R. Straatman, Malaise trap (BISH). Kolombangara I.: 1 9, Pepele, 30 m, 7 Febr., 1964, coll. P. Shanahan, Malaise trap (BISH). Vella Lavella I.: 1 9, Ulo Crater, 10 m, Dec., 1963, Malaise trap (BISH). Bougainville I.: 1 4, Naval Air Base, 25 April, 1945, coll. G. E. Bohart (USNM). Psenulus chariis spec. nov. Female. — Head black; mandibles yellow with dark tips, palpi yellowish, anterior margin of clypeus reddish, scape of antennae yellow, flagellum reddish-yellow. Thorax black with following yellow parts: pronotum above, pronotal tubercles, sides of scutum and two incomplete median lines (Fig. 52), scutellum except depressed lateral parts, metanotum and part of its lateral depressions, two parallel lines on posterior surface of propodeum, sides of propodeum, upper part of anterior plate of mesepisternum, a posteriorly somewhat vague semicircular band on mesopleura and almost complete hypo- epimeral area. Axillae black with small yellowish-red dot. Fore and mid legs including coxae dark yellow, hind legs including coxae, and gaster including petiole, yellowish- red. Veins of wings yellowish-red, subcostal vein slightly darker. Anterior margin of clypeus resembling that of P. interstitialis interstitialis, with two small teeth. Raised part of interantennal carina not broadened dorsally. No distinct transverse carina below antennae, only traces laterally. Frons and vertex shining, almost impunctate, lateral parts of frons slightly raised. Occipital carina ending in hypostomal carina. Third segment of antennae about two and a half times as long as broad at apex, fourth segment nearly twice, segments 8—11 about one and a half times as long as broad at apex, 12th segment about twice as long as broad at base. Scutum finely, sparsely, punctate, also with a few larger punctures, scutellum and metanotum finely, sparsely, punctate. Enclosed area of propodeum depressed, with oblique longitudinal carinae. Upper part of back of propodeum smooth, posterior surface with fine carination which is partly obliquely parallel, sides of propodeum with fine reticulate carination. Longitudinal sulcus of propodeum about as long as upper two thirds of propodeum. Mesopleura finely punctate. Anterior oblique suture indistinctly foveolate, widened upper part smooth. First recurrent vein of fore wings about interstitial. Femora thick, apex of mid tibiae with oblique ridge and a few short reddish thorns. Apex of hind tibiae with three short reddish spines, close together. Petiole about as long as hind tibiae, cylindrical. Pygidial area as in the nominate form of P. interstitialis. Face with appressed silvery pubescence and a number of long erect hairs, rest of body yellowish-grey pubescent. Sides of petiole with long erect hairs. Epicnemial areas below with a round patch of appressed silvery pubescence. Hind margin of fourth and fifth gastral sternites with dense and short, whitish, pubescence. Length about 8,5 mm. Male unknown. J. P. van LitH: Oriental Psenulus 187 Solomon Is.: 3 9 (holotype and 2 paratypes), Choiseul I., Malangona, 30 m, 14 March, 1964, Malaise trap, coll. P. Shanahan (BISH). Structurally this wasp is close to P. interstitialis. It is the second known representative of this group in the Solomon Islands. The extent of the black marking of the scutum resembles that of some of the Philippine species belonging to this group. The wing veins, however, are yellowish-red, as in its near relatives from New Guinea and north- eastern Australia, which have a much more yellow coloured scutum. Psenulus araucarius spec. nov. A single female from Central New Guinea differs structurally so much from P. interstitialis Cameron that it seems justified to describe it as a new species. Scutum with square black mark. Outer two-thirds of lateral depressions of scutellum black; depressions of metanotum yellow. Enclosed area and longitudinal sulcus of propodeum not completely black. Metapleura with black streak. Dorsal side of antennae dark brown, ventral side yellowish-red. Veins of fore wings yellowish, subcostal vein darker. Antennae longer, third segment more than three times as long as broad at apex (in P. interstitialis about two and a half times as long as broad) and fourth and fifth segments about two and a half times as long as broad (in P. interstitialis about twice as long as broad), segments 9—11 nearly twice as long as broad at apex, last segment more than twice as long as broad at base. Distance between clypeal teeth large, nearly one-fifth of total distance there. Pygidial area much broader at base (Fig. 53, araucarius; Fig. 42, holotype of nominate subspecies). Base of hind tibiae dorsally smooth and flattened, this area margined by a horseshoe-shaped line of small blunt thorns. Usually this area is not flat and thorns are irregularly placed. In some P. interstitialis from NE New Guinea the structure of the tibial base resembles that of P. araucarius. As in P. interstitialis, the anterior oblique suture is linear, the widened upper part smooth. Pubescence of face pale golden, mostly appressed but also with long, erect, hairs. Posterior margin of fourth and fifth gastral sternites with dense and short pale pubes- cence. Length about 9 mm. Male unknown. Central N New Guinea: 1 9 (holotype), Sahuweri River Area, Araucaria River Val- ley, Araucaria Camp, 800 m, March, 1939, coll. L. J. Toxopeus, Neth. Ind.-American New Guinea Expedition (ML). The right antenna is missing. This specimen, already labelled by Dr. Karl V. Krombein “new species”, is easily distinguished by the wider pygidial area and by the much longer antennae. Psenulus segrex spec. nov. Female. — Resembling P. interstitialis. Anterior margin of clypeus reddish-brown. Scape and following segment of antennae yellow, flagellum reddish. Scutum yellow, marking resembling that of P. /wzonensis: a large square central black spot, connected by a narrow line with the black triangle on the posterior margin of the scutum; along the parapsidal sutures a narrow, longitudinal, black streak, about as long as the central 188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Fig. 52-56. Species of Oriental Psenulus. 52, thorax of female of P. chariis sp. nov. (Solomon Is.); 53, pygidial area of female of P. araucarius sp. nov. (New Guinea); 54—56, scutum, second and third submarginal cells of left fore wing, pygidial area of female of P. segrex sp. noy. (Celebes) spot (Fig. 54). Scutellum and axillae yellow, lateral depressions black. Metanotum yellow, inner third of lateral depressions black. Propodeum yellow, enclosed area and median longitudinal sulcus black. Pronotum yellow, lower part and a mark against anterior margin of epicnemial areas black. Mesosternum black, marking roundly dilated on to lower part of sides of thorax, which are yellow, including hypo-epimeral and subalar areas and anterior plate of mesepisternum. Hind margin of mesopleura and the whole metapleura black. Fore and mid legs yellow, including coxae, but fore tarsi and mid tibiae and tarsi reddish. Hind legs reddish except coxae which are yellow. Subcostal vein of fore wings brown, other veins and stigma yellow or yellowish-brown. Tegulae yellow. Gaster in- cluding petiole reddish, apex of petiole dorsally slightly brown. Face without transverse carina below antennae. Distance between clypeal teeth less than one-fifth of total distance there between the eyes. Third antennal segment about two and a half times as long as broad at apex, 10th and 11th segments about one and a half times as long as broad at apex, last segment more than twice as long as broad at J. P. van LITH: Oriental Psenulus 189 base. Upper part of propodeum with distinct oblique striae, lower part with more irregular carination. Anterior oblique suture almost linear, widened part smooth. Base of hind tibiae flattened on dorsal side and with horseshoe-shaped row of nine small, dark red teeth; apical margin with three small, dark red, teeth. First recurrent vein of fore wings ending a short distance before the end of the first submarginal cell (Fig. 55). Pygidial area narrow (Fig. 56). Second gastral sternite with semi-circular, not well- defined, depression. Pubescence of face pale golden. Body with yellowish pubescence, posterior margin of fourth and fifth gastral sternites with short and dense whitish pubescence. Length about 10,5 mm. Male unknown. Celebes: 1 9 (holotype), Bonthain, Wawa Karaeng, 1100 m, Sept.—Oct., 1931, coll. G. Heinrich (ZMB). P. segrex and P. lamprus were collected simultaneously, at an altitude of 1100 meters. P. segrex is closer to P. interstitialis Cameron, which also has yellow veins (except the subcostal vein) and stigmata, whilst P. /amprus seems to be more related to the Philip- pine forms of the P. interstitialis group, which have brown veins and stigmata. It seems possible that the two species from Celebes are what L. J. Toxopeus called a “species duplex”, but very few specimens could be studied and more material from this island is needed. Psenulus luzonensis luzonensis (Rohwer) 1921, Rohwer, Phil. Jl. Sc. 18 (3): 313-314, ® (Diodontus luzonensis; Luzon). 1962, Van Lith, Zool. Verh. Leiden 52: 86-87, ® and & (Psenulus interstitialis luzonensis, Luzon, Biliran, Negros). 1970, Van Lith, Steenstrupia 1: 100-101 (Tawi Tawi). New records from the Philippine Islands: Luzon: 1 ® and 3 4, Atimonan, Quezon, July and August, 1908, coll. Micholitz (ZMB); 1 9, Antipolo, Rizal, 6 Sept., 1952, coll. Townes family (Coll. Townes); 1 9, Mt. Isarog, Camarines Sur, 500 m, 4 April, 1963, coll. H. M. Torrevillas (BISH); 1 ©, Cuenca, Batangas, 29 Jan., 1954, coll. Macabasco (BPIM); 1 9, Bigti, Bulucan, 26 Sept., 1961, coll. C. Baltazar (BPIM); 1 &, Mt. Montalban, Wa-Wa Dam, Rizal, 150—200 m, 10 March, 1965, coll. H. M. Torrevillas (BISH); 1 ©, Sierra Madre, Nuevacija, March, 1965, coll. A. Villareal (CNC). Mindanao: 1 9, Parang, Cotabato, 16 June, 1953, coll. Henry Townes (Coll. Townes). First record from Mindanao. There is some variation in the extent of the black marking of the thorax. The female from Mindanao has two small, lateral, comma-shaped, black marks. In the females from Mt. Isarog and from Antipolo the scutum has only one median square spot. In the female from Bigti the median spot on the scutum is reduced to two small, narrowly separated, oblong marks, well between prescutal sutures, if these would be continued. Also the black marking of the mesopleura, ventral side of thorax and of the depressed lateral parts of the scutellum is variable. The pubescence of the clypeus is always silvery; of the frons distinctly pale golden in the Luzon females, pale yellow in the female from Mindanao. 190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Psenulus luzonensis pseudolineatus Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 88, & (Palawan, Philippine Is.). 1970, Van Lith, Steenstrupia 1: 101, & (Balabac, Philippine Is.). New record from Palawan: 1 &, 3 km NE of Tinabog, 12 May, 1962, coll. H. Holt- mann, Malaise trap (BISH). This is the second record of a male belonging to this subspecies. The hind femora are light brown, base not darkened, as in the holotype. A narrow line connects median black mark of scutum with broad black hind margin (Fig. 57). Depressed lateral parts of scutellum black, only the inner part of the depressed lateral areas of metanotum is black. Veins of wings brown. Third antennal segment about two and a half times and segments 4—8 about twice as long as broad at apex, following segments somewhat shorter, last segment longer than twice as broad at base. Segments 8—12 somewhat rounded below. Anterior oblique suture narrow, foveolate, widened upper part smooth. Psenulus luzonensis davanus (Rohwer) 1923, Rohwer, Phil. Jl. Sc. 22 (6): 599, & (Diodontus davanus; Mindanao). 1962, Van Lith, Zool. Verh. Leiden 52: 87-88, 2 and & (Psenulus interstitialis davanus, Min- danao and Mindoro). This subspecies is now also recorded from the Philippine island Balabac: 1 ©, 10 km S of Balabac, 26 May, 1962, coll. H. Holtmann, Malaise trap (BISH). Anterior oblique suture narrow, weakly foveolate, widened upper part smooth and shining. Second gastral sternite without semicircular depression. The dense, appressed pubescence of clypeus and tempora is silvery-white, pubescence of frons pale golden. Psenulus lamprus spec. nov. Two Psenulus females from Celebes resemble P. interstitialis from New Guinea. The thorax is yellow, including lower part of pronotum, but except a black triangle on posterior margin of scutum, black scutellar suture and two small dark spots in the enclosed area of the propodeum. Anterior margin of clypeus reddish. Scape of antennae and following segment yellow, flagellum yellowish-red, only slightly darkened on upper side. Unlike the forms from New Guinea and NE Australia, the veins of the wings are brown, instead of yellowish. All coxae, fore legs, and trochanters, femora and tibiae of mid legs yellow. Mid tarsi and hind legs including trochanters yellowish-red. Gaster including petiole yellowish-red. Distance between teeth on anterior margin of clypeus as in P. interstitialis. Third antennal segment about two and a half times as long as broad at apex, fourth segment twice as long, segments 5—8 about one and a half times, segments 9—11 little longer than they are broad at apex, last segment about twice as long as broad at base. Back of propodeum, and sides except anterior margin, with dense and rather coarse reticulate carination, upper half with parallel oblique striae. Anterior oblique suture narrow, weakly foveolate, widened upper part smooth and shining. Apex of hind tibiae with three small reddish spines. First recurrent vein of fore wings interstitial. Petiole cylin- drical. Pygidial area narrow. J. P. VAN LITH: Oriental Psenulus 191 Pubescence of face and frons very pale golden, almost silvery, rest of body yellowish- grey, posterior margin of fourth and fifth gastral sternites with short and dense, whitish, pubescence. Length about 9 mm. Male unknown. Celebes: 1 ® (holotype), Bonthain, Wawa Karaeng, 1100 m, end Aug., 1931; 1 9 (paratype), Bantimurung, Aug., 1931, both coll. G. Heinrich (holotype ZMB, paratype in the author’s collection). This species is easily distinguished from P. interstitialis from New Guinea and NE Australia by the completely yellow lower part of the pronotum and by the brown veins of the fore wings. This latter character may point to a close relationship with the Philip- pine forms, which all have brown wing veins, but which have more extended black markings on the scutum and rest of the thorax. Psenulus annamensis spec. nov. Female. — Head black; labrum reddish, mandibles yellow with red tips, palpi yellow- ish, scape of antennae yellow, flagellum black-brown above, reddish below. Thorax black; the following parts yellow: pronotum dorsally, pronotal tubercles, a band along lateral margins of scutellum almost reaching anterior margin, two median lines on scutum (Fig. 58), scutellum and metanotum including postero-lateral margins but except a small black mark on posterior margin of scutellum and the black lateral depressions, two large longitudinal marks on posterior face of propodeum, similar marks on the sides, their upper parts vaguely connected with the inner marks, upper third of anterior plate of mesepisternum, a vague spot on the mesopleura behind the anterior oblique suture. Axillae black but with a very small reddish spot in the hind corner. Fore and mid legs yellow including greater part of coxae, tarsi somewhat reddish, hind legs reddish but coxae yellow and trochanters brown, outer spur of hind tibiae yellowish. Petiole reddish, apex darkened, gaster reddish. Tegulae yellow. Veins of wings dark brown. Protruding median part of anterior clypeal margin almost straight, a small tooth on the corners, distance between the teeth less than one-fifth of total distance there between the eyes; disk of clypeus indistinctly punctate. Frons and vertex with minute punctures. Interantennal carina raised, narrow, no transverse carina below antennae. Third antennal segment more than three times, segments 4—6 more than two times as long as broad at apex, following segments gradually shorter, 11th segment about one and a half times as long as broad at apex; last segment more than twice as long as broad at base. Legs normal, apex of mid tibiae somewhat depressed longitudinally, this depression margined anteriorly by an oblique ridge, posteriorly by three red thorns. Apex of hind tibiae with three short red thorns. First recurrent vein of fore wings interstitial, second recurrent vein ending well in third submarginal cell. Petiole long, cylindrical. Gaster with very fine punctures, pygidial area long and narrow, carinae on apical half parallel. Second gastral sternite with indistinct semi-elliptic depression. Face with appressed silvery pubescence and also with long and erect hairs. Rest of body yellowish-grey pubescent, densest on mesosternum. Epicnemial areas below with circular pubescent spot. Hind margin of fourth and fifth gastral sternites and disk of sixth sternite with dense pubescence. 192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Fig. 57—60. Species of Philippine and South Asiatic Psenulus. 57, thorax of male of P. luzonensis pseudolineatus Van Lith (Palawan); 58, thorax of female of P. amnamensis sp. nov. (Vietnam); 59, thorax of female of P. annamensis subsp. ? (India, Sikkim); 60, thorax of female of P. an- namensis subsp. ? (S India) Length about 9,5 mm. Male unknown. Vietnam: 1 9 (holotype), Dai Lanh, N of Nha Trang, 30 Nov.—5 Dec., 1960, coll. C. M. Yoshimoto (BISH). This female is closely related to P. elegans (Van Lith, 1962) from Java, which has been placed in the group of P. interstitialis. However, the axillae of P. elegans are yellow J. P. van LITH: Oriental Psenulus 193 and its scutellum is largely black. Unfortunately the female of the Java form is unknown. It would be interesting to know the length of its antennal segments, as in the forms from the southeastern Asiatic continent the third antennal segment is longer than in the closely related P. petersen? (Van Lith, 1970) from the Philippines and most of the other Indo-Australian species of the group of P. interstitialis. A female from Sikkim is very similar. It differs in the somewhat larger size, the yellow lines on the scutum are narrower (Fig. 59), the yellow spot on the anterior plate of the mesepisternum is smaller, the mesopleura are completely black, the fore side of the hind femora is dark brown and the posterior surface of the hind femora is yellowish. Length about 10,5 mm. India: 1 9, Sikkim, coll. Bingham (ZMB). Another female, from S India, is also very similar and closely related. Scutum and scutellum are slightly darker than in the form from Vietnam (Fig. 60), the marks on posterior surface and sides of propodeum are fused into two large square marks, fore and mid femora have a large brown streak on hind surface, mid tibiae are brown, hind legs dark brown except a yellowish streak on hind surface and paler base of tibiae. Sides of thorax black except a vaguely defined small yellow spot on the upper part of the hypo- epimeral area. Axillae with a very small yellowish spot. I expect that this will prove to be a different subspecies, either of P. e/egans or of P. annamensis. Before definitely naming this Indian form more specimens from this area should be available. S India: 1 9, Nilgiri Hills, Devala, 3200 ft, Oct., 1960, coll. P. Susai Nathan (ML). Psenulus hoozanius spec. nov. Male. — Head black; mandibles yellow except dark tips, palpi straw-yellow, scape of antennae yellow, dorsally with brown streak, second and following antennal segments dorsally dark brown, underside pale yellowish-brown, but underside of last five segments light brown. Thorax black with following yellow parts: dorsal part of pronotum, pronotal tubercles, a narrow mark on scutum (Fig. 61) along the tegulae, two small marks near hind margin of scutum, two irregular marks on scutellum, metanotum except lateral depressions, two parallel elongate marks on posterior side of propodeum. Fore side of fore coxae and complete trochanters pale yellow, femora, tibiae and tarsi darker yellow- ish. Mid trochanters pale yellow, tips of coxae, femora, tibiae and tarsi darker yellowish. Hind coxae yellowish-brown with dark brown triangle, rest of hind legs dark brown, ventrally somewhat paler, tibiae with yellowish base, tarsi dark brown. Veins of wings brown. Petiole pale reddish with darkened apical part and brown ventral plate; rest of gaster reddish, apical spine dark brown. Clypeus convex with coarse coriaceous sculpture including the distinctly depressed margin, which is bidentate. Raised part of interantennal carina narrow dorsally, ending in transverse carina below antennae. Frons and vertex smooth and shining, almost impunctate. Occipital carina ending normally in hypostomal carina. Antennae slender, third segment about two and a half times as long as broad at apex, fourth segment twice, segments 5—9 slightly more than twice, segments 10—12 about twice as long as broad at apex, last segment nearly three times as long as broad at base. No tyloidea. Scutum with fine, wide-spread, puncturation and a few larger punctures. Scutellum and metanotum almost impunctate. Enclosed area of propodeum depressed, with oblique 194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 64 Fig. 61—65. Species of Oriental Psenulus. 61—62, thorax, second and third submarginal cells of right fore wing of male of P. hoozanius sp. nov. (Formosa); 63—65, clypeus, scutum, pygidial area of female of P. ceylonicus sp. nov. (Ceylon) longitudinal carinae. Median longitudinal sulcus of propodeum as long as upper two- thirds of propodeum, upper third of hind surface of propodeum smooth, lower two- thirds of back and greater part of sides of propodeum with irregular carination. Mesopleura finely punctate, metapleura very smooth. Anterior oblique suture foveolate, with long transverse carinae, widened upper part smooth. Legs slender. First recurrent vein of fore wings interstitial, second recurrent vein ending well in third submarginal cell (Fig. 62). Gaster slender, finely punctate, petiole cylindrical. Face with appressed silvery pubescence and a few longer, erect, greyish hairs. Rest of body with greyish pubescence, mesosternum with long whitish pubescence. Length about 8 mm. Female unknown. Formosa: 1 4 (holotype), Hoozan, Oct., 1910, coll. H. Sauter (ZMB). P. hoozanius has been placed provisionally into the group of P. interstitialis. A study of the female necessary to obtain more knowledge about its relationships. In com- parison with the Philippine species of this group the thorax is darker. J. P. van LITH: Oriental Psenulus 195 Psenulus ceylonicus spec. nov. Female. — Head black, mandibles yellow with dark tips, palpi yellowish. Scape of antennae yellow, flagellum dark brown above, pale brown below. Dorsal side of pro- notum and pronotal tubercles yellow. Scutum yellow, two broad black lateral lines reaching pronotum anteriorly, posteriorly connected with black hind margin of scutum, median black line complete (Fig. 64). Tegulae yellowish transparent. Scutellar suture black; scutellum, axillae and metanotum yellow, lateral depressions of scutellum black, inner third of lateral depressions of metanotum black. Propodeum yellow with broad, black, basal band (as broad as enclosed area), black sulcus and narrow black posterior margin. Ventral part of thorax and epicnemial areas black, mesopleura black with yellow spot against lower part of anterior oblique suture; anterior plate of mesepisternum yellow, hypo-epimeral area yellow, subalar area of fore wings black. Metapleura black. Fore and mid legs including coxae yellow. Coxae of hind legs yellow, trochanters brown, femora yellowish-red, dorsal side brown, tibiae reddish-brown but extreme base and apex darkened, hind metatarsus dorsally brown. Veins of wings brown. Basal half of petiole yellowish-red, apical half black. Rest of gaster, including ventral plate of petiole, reddish. Clypeus slightly convex and finely punctate, broad margin depressed, almost impunct- ate and shining, distinctly bidentate, distance between teeth about one-fifth of distance between eyes (Fig. 63). No transverse carina below antennae, interantennal carina sharp, strongly protruding between antennae, frons and vertex very finely punctate. Apex of mandibles bidentate. Antennae somewhat clavate, third segment more than two and a half times as long as broad at apex, segments 4—6 about twice and segments 7—9 about one and a half times as long as broad at apex, segments 10—11 slightly shorter, 12th segment about twice as long as broad at base. Scutum sparsely punctate, prescutal sutures distinct on anterior third of scutum. Enclosed area of propodeum triangular, behind enclosed part a large smooth and shining area. Back of propodeum finely, irregularly, reticulate. Anterior oblique suture foveolate, widened upper part smooth. First recurrent vein of fore wings ending just before second submarginal cell, almost interstitial. Second recurrent vein ending in third submarginal cell. Petiole almost cylindrical. Gaster with extremely fine punctatioon. Pygidial area narrow, broadest at base and middle part slightly narrower than apex (Fig. 65). Pubescence of face silvery, also of frons below ocelli and on temples. Rest of body with whitish pubescence, posterior margin of fourth and fifth gastral sternites with short and dense, whitish, pubescence. Epicnemial areas below with circular pubescent spot. Mesosternum densely pubescent medianly. Length about 9 mm. Male unknown. Ceylon: 2 (holotype and paratype), Kandy, June, 1918, coll. O. S. Wickwar (BM 1919—252). In the paratype the lateral black bands on the scutum are slightly broader than in the holotype; the yellow parts of the paratype are discoloured. P. ceylonicus differs from a similar S Indian form (cf. P. annamensis), in having the thorax more yellow, especially the sides. 196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Psenulus bakeri bakeri (Rohwer) 1921, Rohwer, Phil. Jl. Sc. 18 (3): 312-313, 2 (Diodontus bakeri; Luzon). 1962, Van Lith, Zool. Verh. Leiden 52: 98-99, ® and 3 (Psenulus bakeri bakeri, Luzon, Min- danao, Samar). New records: Luzon: 1 G', Los Baños, Laguna, 29 Nov., 1952, coll. Townnes family (Coll. Townes); 1 9, Mt. Makiling, Laguna, 8 Nov., 1954, coll. C. Baltazar, 1 9, Tiaong, Quezon, 8 Aug., 1957, coll. A. Concepcion, 1 9, Atimonan, Quezon, Aug. 1963, coll. A. Concepcion (BPIM); 1 9, Abatan, Buguias, Mountain Prov., 60 km S of Bontoc, 1800—2000 m, 9 June, 1964, coll. H. M. Torrevillas (BISH). Mindanao: 1 8, Agusan, Los Arcos, 19—23 Nov., 1959, Malaise trap, coll. L. Quate and C. Yoshimoto (BISH). In the male from Los Baños the first recurrent vein of the fore wings ends distinctly in the first cubital cell. This proves again that the course of the recurrent veins may be variable and therefore is not a reliable character for identification. The anterior oblique suture is nearly simple, the widened upper part is smooth and shining. Group of Psenulus pulcherrimus Psenulus pulcherrimus projectus Van Lith 1962, Van Lith, Zool. Verh. Leiden 52: 102-103 (Java, Krakatau). 1969, Van Lith, Tijdschr. v. Ent. 112: 200 (Kangean I.). New record: 1 9, W Java, Tjipanas, 5—10.000 ft, 8 Sept., 1965, coll. J. E. Lukavsky (CNC). Psenulus xanthognathus xanthognathus Rohwer 1910, Rohwer, Proc. U.S. Nat. Mus. 37: 660, 4 (Psenulus (Neofoxia) xanthognathus, Luzon). 1962, Van Lith, Zool. Verh. Leiden 52: 104-107, partim (Psenulus carinifrons xanthognathus). 1966, Van Lith, Tijdschr. v. Ent. 109: 44-45, partim (Psenulus carinifrons xanthognathus). 1969, Van Lith, Tijdschr. v. Ent. 112: 206 (Psenulus xanthognathus xanthognathus). New record: 1 3, Luzon, Sierra Madre, Nuevacija, March, 1965, coll. A. Villareal (CNC). Psenulus xanthognathus centralis Van Lith 1969, Van Lith, Tijdschr. v. Ent. 112: 207-208 (SE Luzon, Masbate, Leyte, Panay, Busuanga). Culion: 1 9, 6 km W of Culion, 6—7 June, 1962, low vegetation, coll. H. Holtmann (BISH). This is the first record from the Philippine island Culion. From Busuanga I., north of Culion, four males, no females, are known. Basal half of fore and mid femora black-brown, as in form “B” from Palawan and Mindanao, but yellow marking on pronotum, scutellum and metanotum not reduced. Apical third of hind tibiae, when seen from aside, black. J. P. van LITH: Oriental Psenulus 197 LIST OF PALAEARCTIC, INDO-AUSTRALIAN AND W PACIFIC Psenulus SPECIES In this list also the Palaearctic species have been divided tentatively into groups. The following species have not been included, as their systematic place is still unclear: P. tathorinus Strand, 1915 — Formosa P. clavatus (Cameron, 1890) — India P. montanus (Cameron, 1907) — India P. puncticeps (Cameron, 1907) | — India I have not yet examined the latter three species; P. montanus is probably very close to P. atratus, P. puncticeps may be near to P. antennatus. Genus Psenulus Kohl Group of fuscipennis fuscipennis (Dahlbom, 1845), 9 & — Europe, Turkey, Siberia, Korea, fuscipennis japonicus Tsuneki, 1959, 9 3 Japan atratus (Fabr., 1804), © d — Europe, Turkey, Morocco, Syria, Siberia atratus puncticeps Gussakovskij, 1932, 9 3 — Europe, Siberia, China, Korea, Japan atratus yamatonis Tsuneki, 1959, © 3 — Japan brevitarsis Merisuo, 1937, & 2 — Europe laevis Gussakovskij, 1930, 9d — Asia meridionalis de Beaumont, 1961, 9 & — S Europe, Crete, Turkey, Irak pan de Beaumont, 1967, 9 g' — SW Asia (Turkey) tanakai Tsuneki, 1959, 9 gd — Japan bisicatus spec. nov, 9 d' — W Pakistan Group of berlandi berlandi de Beaumont, 1937, 9 — Europe Group of concolor concolor (Dahlbom, 1845), 2 & — Europe, Turkey, Japan schencki Tournier, 1889, 9 gd — Europe, Turkey lubricus (Pérez, 1905), 9 oS — Japan maculipes Tsuneki, 1959, 9 d' — Japan nipponensis Yasumatsu, 1942, 9 — Korea, Japan Group of /aevigatus laevigatus Schenck, 1857, 9 3 — Europe fuscipes Tsuneki, 1959, 9 3 — Japan Group of nikkoensis nikkoensis Tsuneki, 1959, 9 4 — Japan 198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Group of anomoneurae (genus Nipponopsen Yasumatsu, 1938) anomoneurae (Yasumatsu, 1938), ® g' — Japan (syn. Psenulus mandibularis Tsuneki, 1959) Group of suifuensis suifuensis spec. nov., 9 — China Group of formosicola formosicola Strand, 1915, 9 & — Formosa Group of compactus compactus Van Lith, 1962, 9 — Sumatra Group of rufobalteatus rufobalteatus Cameron, 1904, 9 — N India (Assam) bicinctus Turner, 1912, © & — N India (Assam, Sikkim), Nepal Group of scutatus scutatus scutatus (Rohwer, 1921), 9 & — Luzon, Samar, Negros scutatus sibuyanensis Van Lith, 1962, 8 — Sibuyan scutatus mindanaoensis (Rohwer, 1923), © — Mindanao scutatus borneensis (Rohwer, 1923), 9 — Borneo hemicyclius Van Lith, 1962, 9 8 — Palawan baltazarae baltazarae Van Lith, 1962, 9 — Luzon baltazarae luteus Van Lith, 1962, 9 — Sibuyan maai Van Lith, 1967, 8 — Borneo dentatus Van Lith, 1962, 9 — Java Group of rugosus rugosus Van Lith, 1962, 9 — Mindanao yoshimotoi yoshimotoi Van Lith, 1969, 9 — Borneo yoshimotoi pontilis Van Lith, 1970, 9 — Tawi Tawi Group of quadridentatus quadridentatus quadridentatus Van Lith, 1962, — Malaya, Vietnam Ld quadridentatus formosanus Tsuneki, 1966, 3 — Formosa separatus Van Lith, 1962, 9 — Java, Borneo pendleburyi Van Lith, 1962, 9 — Borneo macrodentatus Van Lith, 1962, 9 — Sumatra parvidentatus spec. nov., 9 — Formosa Group of nasicornis nasicornis spec. nov., 9 — Celebes J. P. van LITH: Oriental Psenulus 199 Group of antennatus antennatus (Rohwer, 1923), © d' — Malaya, Singapore, Java, Bali penangensis (Rohwer, 1923), 9 — Malaya mauritii Van Lith, 1969, 9 — Malaya bengalensis spec. nov, G' — India (Bengal) nietneri spec. nov, 9 — Ceylon noonadanius Van Lith, 1970, 9 & — Balabac, Palawan Group of singularis singularis Van Lith, 1962, 2 — Luzon Group of erraticus erraticus erraticus (Smith, 1861), ® — Celebes erraticus basilanensis (Rohwer, 1921), 9 & — Singapore, Vietnam, Borneo, Tawi Tawi, Basilan erraticus butuanensis Van Lith, 1962, 5 — Mindanao Group of crabroniformis crabroniformis crabroniformis (Smith, 1858), — Borneo, Thailand Ce crabroniformis sumatranus (Ritsema, 1880), @ — Sumatra, Java crabroniformis subsp. ?, 9 — Busuanga philippinensis philippinensis (Rohwer, 1921), 9 — Luzon philippinensis dapitanensis (Rohwer, 1923), ® | — Mindanao, Bohol Group of tuberculifrons tuberculifrons tuberculifrons (Rohwer, 1921), ® — Luzon tuberculifrons decoratus Van Lith, 1962, 2 — Sibuyan maurus (Rohwer, 1921), 3 — Luzon suluensis Van Lith, 1970, 9 & — Tawi Tawi cavifrons Van Lith, 1962, 2 — Samar, Mindanao maculatus maculatus Van Lith, 1962, 9 — Malaya maculatus javanensis Van Lith, 1962, 9 G' — Java Group of nigrolineatus nigrolineatus nigrolineatus (Cameron, 1907), — Malaya, Borneo Ne nigrolineatus ajax (Rohwer, 1921), ® — Luzon nigrolineatus flavicornis Van Lith, 1962, 9 — Sibuyan nigrolineatus dubius Van Lith, 1962, 9 — Mindanao, ? Tawi Tawi nigrolineatus sulphurens Van Lith, 1962, 9 — Sumatra diversus Van Lith, 1962, 9 — Singapore, Vietnam Group of /uteopictus luteopictus luteopictus (Rohwer, 1921), 9 d — Luzon, Negros luteopictus calapanensis Van Lith, 1962, 9 — Mindoro 200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 Group of sandakanensis sandakanensis (Rohwer, 1923), Q & — Borneo, ? Sumatra filicornis (Rohwer, 1923), dg — Basilan ajaxellus (Rohwer, 1923), 8 — Mindanao trimaculatus Van Lith, 1962, 3 — Malaya pagdeni Van Lith, 1962, 9 — Malaya, Borneo Group of esuchus esuchus (Rohwer, 1923), 9 gd — Borneo, Balabac corporaali Van Lith, 1962, 9 — Sumatra varius Van Lith, 1962, 2 d — Malaya pallidicollis spec. nov., 9 — Borneo neptunus spec. nov., 2 — Celebes Group of pseudajax pseudajax pseudajax Van Lith, 1962, 4 — Java psendajax holtmanni subspec. nov, 5 — Culion, Busuanga Group of interstitialis interstitialis interstitialis Cameron, 1906, 9 & — New Guinea interstitialis lutescens (Turner, 1907), 9 — NE Australia interstitialis salomonensis subspec. nov., © & — Solomon Is. chariis spec. nov., 9 — Solomon Is. arancarius spec. nov., 9 — New Guinea segrex spec. nov., 9 — Celebes luzonensis luzonensis (Rohwer, 1921), 9 & — Luzon, Biliran, Negros, Tawi Tawi, Mindanao luzonensis pseudolineatus Van Lith, 1962, 3 — Palawan, Balabac luzonensis davanus (Rohwer, 1923), 9 dg — Mindanao, Mindoro, Balabac luzonensis baliensis Van Lith, 1962, 9 — Bali nigromaculatus (Cameron, 1907), 9 — Borneo multipictus (Rohwer, 1921), 9 — Luzon tristis Van Lith, 1962, & — Palawan ornatus (Ritsema, 1876), & — Java elegans Van Lith, 1962, 3 — Java lamprus spec. nov., 9 — Celebes peterseni Van Lith, 1970, 9 — Tawi Tawi continentis Van Lith, 1962, 9 — Malaya annamensis spec. nov., 9 — Vietnam hoozanius spec. nov., d' — Formosa ceylonicus spec. nov., 9 — Ceylon bakeri bakeri (Rohwer, 1921), 9 & — Luzon, Mindanao, Samar bakeri boholensis Van Lith, 1962, 9 — Bohol bakeri canlaonensis Van Lith, 1962, 9 — Negros J. P. van LITH: Oriental Psenulus 201 Group of pulcherrimus pulcherrimus pulcherrimus (Bingham, 1896), Xf pulcherrimus eburneus Van Lith, 1969, 9 pulcherrimus projectus Van Lith, 1962, 9 & xanthonotus Van Lith, 1969, 9 melanonotus Van Lith, 1969, 8 carinifrons carinifrons (Cameron, 1902), & carinifrons scutellatus Turner, 1912, 9 & carinifrons rohweri Van Lith, 1962, 9 gd carinifrons malayanus Van Lith, 1969, 9 & carinifrons bismarckensis Van Lith, 1970, 9 carinifrons subsp. ?, q extremus Van Lith, 1966, 9 & sinclairi Lal, 1939, 3 sogatophagus Pagden, 1933, 9 dg xanthognathus xanthognathus (Rohwer, 1910), Digi xanthognathus centralis Van Lith, 1969, £ & xanthognathus centralis form A, © G' xanthognathus centralis form B, 9 gd iwatai Gussakovskij, 1934, © dg (subgenus Eopsenulus Gussakovskij, 1934) REFERENCES Tenasserim, Vietnam N India (Bengal) Java, Krakatau, Kangean Is. Formosa Sumbawa N India NE Australia, Papua, Ambon, Buru, Mindanao Java, Kangean Is., Luzon, Formosa Malaya, Singapore, Vietnam, Sumatra, Borneo Bismarck Is. S India New Guinea India (Bombay) Malaya, Thailand, ? India (Assam) Luzon SE Luzon, Masbate, Samar, Leyte, Cebu, Panay, Busuanga, Balabac, Culion Negros, Sibuyan Palawan, Mindanao Japan Cameron, P., 1890. Hymenoptera orientalia, or contributions to a knowledge of the Hymenoptera of the Oriental Zoological Region. — Mem. Proc. Manch. Lit. Phil. Soc. [4] 3: 239-284. India. — Ann. Mag. Nat. Hist. [7] 13: 211-233. Tijdschr. v. Ent. 49: 215-233. 1904. Descriptions of new species of Aculeate and Parasitic Hymenoptera from Northern , 1906. Hymenoptera of the Dutch Expedition to New Guinea in 1904 and 1905, Part I. — 1907. Description of New Species of Hymenoptera from Borneo. On two new species of Mellinus. — Jl. Straits Br. Asiat. Soc. 48: 22-25. Gussakovskij, V., 1934. Beitrag zur Kenntnis der Pseninen- und Pemphredoninen-Fauna Japans (Hy- menoptera, Sphecidae). — Mushi 7 (2): 79-89. Handlirsch, A., 1887. Monographie der mit Nysson und Bembex verwandten Grabwespen, II. — Sitzb. kais. Akad. Wissensch. 96: 219-311. Lith, J. P. van, 1962. Contribution to the knowledge of the Indo-Australian Pseninae (Hymeno- ptera, Sphecidae), Part II. Psenulus Kohl, 1896. — Zool. Verh. Leiden 52: 1-118. —-—, 1965. Contribution to the knowledge of the Indo-Australian Psenini. Part III. New species of the subgenera Psen Latreille and Mimumesa Malloch and a review of East-Asiatic and Indo-Australian Psen Latreille, s.l. — Zool. Verh. Leiden 73: 1-80. > TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 3, 1972 1966. The group of Psenulus pulcherrimus (Hymenoptera, Sphecidae). — Tijdschr. v. Ent. 109: 35-48. 1967. Contribution to the knowledge of the Bornean Psenini (Hymenoptera, Sphecidae). — Ent. Ber. Amst. 27: 15-20. 1969. Descriptions of some Indo-Australian Psenulus and revision of the group of Psenulus pulcherrimus (Bingham) (Hymenoptera, Sphecidae, Psenini). — Tijdschr. v. Ent. 112: 197-212. 1970. The Psenini collected by the Noona Dan Expedition in the Philippine and Bismarck Islands (Insecta, Hymenoptera, Sphecidae). — Steenstrupia 1: 91-105. Rohwer, S. A., 1910. Some new Hymenopterous Insects from the Philippine Islands. — Proc. U.S. Nat. Mus. 37: 657-660. 1921. Descriptions of new Philippine wasps of the subfamily Pseninae. — Phil. Jl. Sc. 18: 309-323. 1923. New Malayan wasps of the subfamily Pseninae. — Phil. Jl. Sc. 22: 593-601. Smith, F., 1858. Catalogue of the Hymenopterous Insects collected at Sarawak, Borneo, Mount Ophir, Malacca and at Singapore, by Mr. A. R. Wallace. — Jl. Proc. Linn. Soc. Zool. 2: 43-130. Strand, E., 1915. H. Sauter's Formosa-Ausbeute. Crabronidae und Scoliidae. III. (Die Gattung Tsuneki, Psenulus nebst Nachtrag zu Sphex.). — Arch. f. Naturgesch., Abt. A 7, 81: 19-23. K., 1959. Contributions to the knowledge of the Cleptinae and Pseninae Faunae of Japan and Korea (Hymenoptera, Chrysididae and Sphecidae). — Mem. Fac. Lib. Arts, Fukui Univ. (2, Nat. Sc.) 9: 1-78. 1966. Contribution to the knowledge of the Pemphredoninae Fauna of Formosa and the Ryukyus (Hymenoptera, Sphecidae). — Etizenia 14: 1-21. 1969. The male of Psenulus tanakai Tsuneki. — The Life Study (Fukui) 13 (3-4): 62. Turner, R. E., 1907. New Species of Sphegidae from Australia. — Ann. Mag. Nat. Hist. [7] 19: 268-276. 1908. Notes on the Australian Fossorial wasps of the Family Sphegidae, with descriptions of new species. — Proc. Zool. Soc. London 30: 463. 1912. Notes on Fossorial Hymenoptera. — X. On new species from the Oriental and Ethiopian regions. — Ann. Mag. Nat. Hist. [8] 10: 361-377. 1916. Notes on fossorial Hymenoptera. — XIX. On new species from Australia. — Ann. Mag. Nat. Hist. [8] 17: 116-136. Yasumatsu, K., 1938. Two new wasps from Japan (Eumenidae and Pemphredonidae). — Mushi 11: 83-86. J. P. van LITH: Oriental Psenulus INDEX The names of new species and subspecies are marked with an asterisk. ajax anomoneurae * annamensis antennatus * araucarius atratus bakeri baltazarae basilanensis * bengalensis bicinctus * bisicatus carinifrons cavifrons centralis * ceylonicus * chariis clavatus concolor corporaali crabroniformis davanus Diodontus diversus elegans Eopsenulus erraticus esuchus formosanus * holtmanni * hoozanius interstitialis iwatai laevis * lamprus luteopictus lutescens luteus luzonensis macrodentatus mandibularis Mellinus meridionalis montanus * nasicornis Neofoxia * neptunus * nietneri nigrolineatus Nipponopsen pagdeni * pallidicollis pan parvidentatus penangensis peterseni projectus Psen pseudajax pseudolineatus pulcherrimus puncticeps quadridentatus rufiventris rufoannulatus rufobalteatus salomonensis sandakanensis scutatus segrex separatus suifuensis suluensis sumatranus taihorinus tuberculifrons varius xanthognathus (The names mentioned only in the review of species occurring in the Palaearctic, Indo-Australian and W Pacific regions have not been included in this index). Postscript. After sending the present article to the printers’, I received Professor K. Tsuneki’s paper “Studies on the Formosan Sphecidae (XIII). A Supplement to the sub- family Pemphredoninae (Hym.) with a key to the Formosan species”, Etizenia 57: 1—21, dated October 25, 1971. Tsuneki considers not only kankauensis Strand, 1915, from Formosa, a subspecies of P. ornatus Ritsema, but also P. tristis Van Lith, from Palawan, and P. elegans Van Lith, from Java. He further describes a new subspecies from Formosa, P. ornatus pempuchten- sis. Unfortunately of all these forms the females, which often show more differentation, are unknown. tn een » En RENT) = Ce Herten nds mn A : ‘ Si No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. 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TIJDSCHRIFT VOOR ENTOMOLOGIE Redactie (Editorial Board) . . . . . . A. Diakonoff, P. J. van Helsdingen, C. A. W. Jeekel, R. de Jong, J. Krikken, M. A. Lieftinck. Address . … . … . . . + . + «Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden. The Journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. It appears in separate issues, forming an annual volume of 350—400 pages. Subscription rate: D.Fl. 62.50 per volume. MONOGRAFIEEN VAN DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING Redactie (Editorial Board) and address as for Tijdschrift voor Entomologie. The Monographs are intended for the publication of larger entomological papers on a single subject and will appear irregularly. The following Monographs have been published: Hors série: F. T. Valck Lucassen et al., 1961. — Monographie du genre Lomapiera Gory & Percheron (Coleoptera, Cetoniidae), 299 pages, 739 figs, 2 pl., map. D.FI. 60.—. No. 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. DIFFERENTIAL USE OF THE HABITAT BY PARDOSA PULLATA (CLERCK) AND PARDOSA PRATIVAGA (L. KOCH) IN A MIXED POPULATION (ARANEAE, LYCOSIDAE) by J. DEN HOLLANDER & H. LOF Department of Biology, Free University, Amsterdam ABSTRACT The distribution of P. prativaga and P. pullata in a mixed population is studied in relation to structural characteristics of the vegetation. The results are discussed in the context of the pos- sibility of the occurrence of interbreeding between the species studied. ACKNOWLEDGEMENTS The authors wish to thank Prof. Dr. L. Vlijm for his valuable criticism during the investigation and the preparation of the manuscript. They are also highly indebted to all the workers of the Biologisch Station “Weevers’ Duin”, Oostvoorne and the ‘‘Ver- eniging tot Behoud van Natuurmonumenten in Nederland’, which made it possible to undertake this study; to Mr. G. W. H. van den Berg, for the drawing of the graphs, and to Miss P. G. Brink, for the typing out of the manuscript. INTRODUCTION The systematics of the species of the Pardosa pullata group are rather confusing. Several authors have described a number of species, under various names, within this group (Locket & Millidge, 1951; Wiebes, 1959; Tongiorgi, 1966; Holm & Kronestedt, 1970). The characters, on which the systematics of the Lycosidae generally are based, almost exclusively concern external morphological features, viz., the colour and design of the body, the spinosity of the legs and the shape of the external genitalia. However, the frequency distributions of a number of measurements of these characters in species of the Pardosa pullata group overlap considerably. Several of the measured characters vary to a rather large extent between populations of the same species; especially the data of mixed populations differ from those of pure populations (Den Hollander, 1970). On the basis of these results, as well as of the occurrence of intermediate specimens in mixed populations, the possibility of interbreeding in the field has been assumed (Locket & Millidge, 1951; Den Hollander, 1970). A study of the phenology of the species Pardosa pullata, P. prativaga and P. sphagnicola has shown that seasonal barriers generally prevent interbreeding in mixed populations, but that in some years, because of weather conditions during spring, these barriers are broken down (Den 205 206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 4, 1972 Hollander, 1971). In addition, under laboratory conditions hybrid specimens have been obtained from interspecific breedings between P. pullata, P. prativaga, and P. sphag- nicola (Den Hollander, in prep.). However, the occurrence of interbreeding in the field is also dependent on the oc- currence of habitat isolation between the species. When two species, occurring in mixed populations, use different parts of the habitat, they will not meet and thus interbreeding will be prevented. Richter et al. (1971) studied the locomotory activity of both P. pullata and P. prativaga in pure, as well as in mixed populations during the reproductive period. They found that this activity of P. prativaga males was much higher than that of P. pullata; the locomotory activity of females was of the same level in both species, this level being considerably lower than in the males. Richter (1970) studied the relation between the structure of the habitat in which the species occurs, the quantity of silk produced, the orientation of the plane in which the silk is spun, and the degree of development of the glandulae ampullaceae. He found that species living in a field-type vegetation (e.g. Pardosa prativaga) have large glandulae ampul- laceae, produce larger quantities of silk, and spin proportionally more silk in the vertical plane, while the converse is true for species from the open ground-type vegetation (e.g. P. pullata). These results suggest a differential use of the habitat by P. prativaga and P. pullata. The aim of the present study is to investigate the distributions of P. pullata and P. prativaga in a mixed population in relation to structural characteristics of the vegetation, in order to establish the differences between the species in more detail. At the same time this study is undertaken with a view to discussing the fore-mentioned differences in relation to the possibility of interbreeding of the species (cf. Den Hollander, 1970, 1971). MATERIAL AND METHODS The study area is a yearly mown meadow, primarily grown with Lys/machia vulgaris L., Hydrocotyle vulgaris L., moss carpets (Polytrichum spec.) and various grasses and sedges (Table 1). It is situated in a dune valley near Rockanje in the southwestern part of the Netherlands. In autumn and winter the field is very wet. It appeared that the water content of the soil was about 50% throughout the study area. However, at the Table 1. The percentages cover of the occurring plant species in the high growing herb layer (K1) and the low growing herb layer (K2) of the vegetation in the study area Plant species Percentage cover High growing herb Lysimachia vulgaris 65 layer (K1) Eupatorium cannabinum 10 Lycopus europaeus 5 Salix repens 10 Scutellaria galericulata 0—5 Low growing herb Hydrocotyle vulgaris 60 layer (K2) Rubus spec. 30 Calamagrostis spec., Carex arenaria, C. nigra, C. disticha 30 Potentilla anserina, P. erecta, P. reptans 10 Polytrichum spec. 0—100 Litter 0—50 J. DEN HOLLANDER & H. LoF: Use of habitat by Pardosa 207 borders, where the ground level was about 30—50 cm higher than in the centre, the water content was about 25% (data of May, 1970). Throughout the study area 68 pitfall traps (diameter 10 cm) were placed (Fig. 1), in such a way that the upper margin of the jars was just level with the surface of the soil. The jars were filled with formalin (4%) to which some detergent was added. In these traps spiders were caught on three days (during 24 hours), between May 28 and June 8. In the same period 10 catches by hand were carried out on five days, two catches of 80 minutes each day. The structural characteristics of the vegetation were described on the basis of the percentage cover, the total height and the species composition (Tables 5 and 6). Two types of herb layer were distinguished, viz., (Table 1) a low growing herb layer and a high growing herb layer. The high layer mainly consisted of Lysimachia, in the low growing type Hydrocotyle was the dominant plant species. In this study, litter is considered a component of the low growing herb layer, so that the complement of the 120m Etra : ' A ' i 8 I isnt ! ak r en i = iI Lamp u: iat Fa ee i Pot A | e en. è e MER + H % è Ÿ i i - PV i i - : 1 I SEE : I A MELO pg pp ee Fig. 1. The location of the 68 pitfall traps in the study area; correspond to locality 1, 2, 3, 4 and 5 of Table 4, en ely) DITO NEO 208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 4, 1972 percentages cover consists of bare ground. With the help of these characteristics four types of differently structured vegetation will be shown to occur in the study area: vegetation of a low open type (LO), a low dense type (LD), a high open type (HO) and a high dense type (HD). The numbers of spiders (males, females and females carrying egg sacs) per pitfall trap in the respective types of vegetation will be compared. The numbers of juveniles were not taken into account, because only small numbers were caught. RESULTS For both species the total numbers per category caught in the pitfall traps over the whole study area are presented in Table 2. It shows that most of the spiders collected were males (about 80%). P. prativaga was caught in the traps slightly more frequently than P. pullata. From the females of P. pullata 44% were carrying an egg sac, whereas this percentage amounted to 28% in P. prativaga. The numbers of spiders (per category) caught by hand are presented in Table 3, showing that less than 20% were males (traps: 80%), largest numbers being females carrying an egg sac (prativaga : 58%; pullata : 80%). Whereas in pitfall traps both species were caught in about equal numbers (Table 2), with hand sampling the num- bers of P. prativaga amounted to less than 30% of the total number. Thus rather striking differences exist between the composition of the sample obtained by hand sampling (Table 3) and that by pitfall trapping (Table 2). Table 2. The total numbers of both species per category caught in 68 pitfall traps over the whole study area (Qc stands for ® with egg sac) Species Category | Jo & of total | Yo ®c of total 9 4 Q Qc total pullata 280 49 39 368 76 44 prativaga 414 63 25 502 82 28 Total: 870, % prativaga of total: 58 These differences illustrate that the two methods of sampling give different results as to the composition of the population. With hand sampling spiders seen by the catcher, i.e. those sitting or walking slowly upon a not too high vegetation, are caught. On the other hand, spiders running on the ground or low in the vegetation are more easily caught in pitfall traps: the higher the locomotory activity, the higher the chance to be caught. Thus, when comparing Tables 2 and 3, it appears that males show the highest locomotory activity of all categories, whereas females carrying an egg sac are the least Table 3. The total numbers of both species per category caught by hand over the whole study area (®c stands for 9 with egg sac) Species Category % & of total % Qc of total 9 ö g Qc total pullata 92 94 380 566 16 80 prativaga 38 72 100 210 18 58 Total: 776, % prativaga of total: 27 J. DEN HOLLANDER & H. Lor: Use of habitat by Pardosa 209 active. In addition, females carrying an egg sac more often occur upon the herb layer than males. The different percentages of P. prativaga in both samples are more difficult to explain, since the real densities of both species are not known. However, Richter et al. (1971) found that, during short periods, the straight line distance covered by P. prativara males was about 3.5 times as long as that of P. pullata males (these data were collected in the present study area). Thus, other factors being equal, the chance that a male of P. prativaga is caught in a trap will be about 3.5 times as great as for a P. pullata male. On the basis of the figures in Table 2 the ratio of the number of males of P. pullata to that of P. prativaga is 2.4 (280: (414: 3.5) = 2.4). As exactly the same proportion is found in hand sampling (Table 2, 92: 38 = 2.4) it may be concluded that males of both species use the habitat in the same way; in other words the chance to be caught is a function of the locomotory behaviour rather than of the way in which the habitat is used. The distributions of both species throughout the study area as established with the help of pitfall trapping are presented in Table 4. The distribution of the traps in the study area is given in Fig. 1. Table 4 shows that both species occur throughout the meadow but that they scarcely occur in locality 1, i.e. the surrounding shrubs and trees. For both species the numbers of specimens are largest in locality 5. In the localities 2—5 the percentage of males of the total catch amounts to the same. Thus both P. prativaga and P. pullata occur especially in the northeastern part of the meadow (see Fig. 1), but the percentage of P. prativaga slightly decreases from locality 2 towards locality 5. The reason may be that towards the northeastern part of the meadow the proportion of the vegetation type LO (see below) increases. Table 4. The distributions of both species per category throughout the study area: , locality 1; — — — , locality 2; ..., locality 3; ——., locality 4; — —., locality 5 (cf. Fig. 1). Locali ( 4 Number Numbers of spiders per 10 traps Wes of traps Fig. 1) prativaga pullata 8 9 total % 4 of} & ® totil % d of | % prativaga of total total total spiders 1 9 6 — 6 100 6 — 6 100 50 2 18 58 11 69 84 25 10 35 nul 66 3 16 68 17 85 80 49 14 63 78 58 4 15 64 16 80 80 44 19 63 70 56 5 10 94 17 111 85 78 19 97 80 54 The results suggest that the factors determining distributions of both P. prativaga and P. pullata are essentially the same. If this is true, a positive correlation of the numbers of P. prativaga and those of P. pullata per trap may be expected. The linear correlation coefficient (r) appeared to be 0.44, which is statistically significant at a = 0,01 (n = 68). Thus, when in one of the 68 traps, i.e. on a certain place in the study area, larger or smaller numbers of P. prativaga occur than in another trap, the same is true for P. pullata. 210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 4, 1972 The percentages cover of the high growing (K1) and low growing (K2) herb layer, as well as the total height of the vegetation (H) for the four vegetation types (cf. Methods), are presented in Table 5. It is evident that in type LO the vegetation is poorly developed, predominantly with low grown plants. Vegetation type LD differs from LO in that, especially, the cover of the low growing herb layer is higher. In type HO the vegetation grows higher than in LD and LO, the percentage cover of the high growing herb layer being slightly higher, that of the low growing herb layer slightly lower, as compared to vegetation type LD. Thus the influence of sunshine and wind-velocity on the microclimate will be less in vegetation type HO than in LO and LD. In type HD the vegetation is densely structured: Table 5. Percentages cover of the high growing herb layer (K1), the low growing herb layer (K2) and the total height of the vegetation in the four types Type % K1 % K2 H LO 8 ( 2—20) 36 ( 5—50) 20 (15—25) LD 24 (15—25) 74 (70—80) 25 (20—30) HO 31 (25—40) 60 (55—65) 37 (30—50) HD 82 (75—95) 81 (70—90) 41 (30—50) both the low growing and the high growing herb layers cover the ground to a large extent. Thus the microclimate will be strongly stabilized against the influence of sunshine and wind-velocity. The total height of the vegetation consequently separates the low types (LO and LD) from the high types (HO and HD). Within the low types there is a separation on the account of the low growing herb layer, in the high types, on that of the high growing. The mean values for the percentages of cover of several plant species in the respective vegetation types are presented in Table 6. The values were obtained in the following manner. First, the vegetation type in the direct surroundings of every pitfall trap was established. Next, the percentages cover of the respective plant species in each of the 68 localities was valued in a range of 0—10 (for moss only the presence or absence was recorded, indicated by 1 or 0, respectively). Table 6 shows that in the low vegetation types (LO and LD) Hydrocotyle is the predominant plant species, while in the high vegetation types (HO and HD) Lysimachia predominates. In the open types (LO and HO) the litter component is larger than in the densely structured types (LD and HD), whereas in the latter the grass component is larger than in the former. Thus, in type LO Hydrocotyle and litter are the most important components. In type LD, as compared with type LO, the components Lys- machia and moss have higher values, the component litter being considerably smaller. The component Lysimachia again is larger in the types HO and HD. In type HD the component grass is larger and the component Hydrocotyle smaller than in all other types. Thus it may be concluded that the four vegetation types differ both in structure (Table 5) and species composition (Table 6). The results of some measurements of microclimatological characteristics (temperature, humidity) are illustrated in Fig. 2. The values presented were taken with an Asmann psychrometer during bright, sunny weather. Fig. 2 shows that at a certain temperature of the air at a height of 2.00 m, the temperature at the ground level is highest in type LO, while the lowest temperature is J. DEN HOLLANDER & H. Lor: Use of habitat by Pardosa 211 ! f ' ' i ' I I I I ' ! 1 ' I N ? ' N LI 1 ; ? 4 40 60 80 100% 22 24 26 28 30 °C RH if Fig. 2. Microclimatological characteristics (temperature, T and humidity, RH) in the various types of vegetation (above: high types; below: low types) recorded in type HD. The differences between air temperature and ground temperature in the high vegetation types is larger when the air temperature is higher. In the low vegetation types these differences are smaller. In all types of vegetation, the relative humidity appears to be highest at the ground level, the value being slightly higher in the densely structured vegetations, as compared with the open structured ones. Especially in the high vegetations (HO, HD) the dif- ference between the relative humidity at the 2.00 m level and at the ground level is higher, when the relative humidity at the 2.00 m level is lower. It may be concluded that the structure of the vegetation exerts an influence on the microclimate, which therefore differs between the respective types (LO, LD, HO and HD). Especially when wind velocity and clouding become important factors, they will modify the microclimate. The above mentioned differences will then become more important. The mean numbers of spiders per trap caught in each of the vegetation types are shown in Table 7. The largest numbers, both of P. prativaga and P. pullata, were caught 212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 4, 1972 Table 6. The mean values for the cover of several plant species in the different types of vegetation (cf. text) Type Plant species Number of traps Lysimachia Hydrocotyle Grasses and Moss Litter Sedges LO 2.0 4.0 2.0 0.3 2EZ 13 LD 4.3 4.9 2.7 0.6 WSS 20 HO 5.6 4.0 2.0 0.2 2.0 11 HD 3.3 342 323 0.0 15 15 in the dense vegetation types (LD and HD), the numbers in HD being larger than in LD. In both types the percentage of P. prativaga in the samples amounts to about 60 %. In the open vegetation types (LO and HO) the total number of spiders is the same, the numbers of P. prativara, however, are lower in LO than in the other three vegetation types, whereas the numbers of P. pullata in LO are the highest, except those in HD. This is reflected in the percentage of P. prativaga in the samples, which also is about 60 % in type HO (as in LD and HD) but only 40 % in vegetation type LO. Throughout the series LO, HO, LD and HD the percentages of the cover of the low growing herb layer increase (Table 5), as do the values for the cover of grasses and sedges, whereas those of the litter decrease throughout the series (Table 6). The numbers of P. prativaga increase gradually from LO through HO and LD towards HD (Table 7), ie. the numbers of P. prativaga are correlated with the cover of the low growing herb layer, grasses, sedges and litter. The same is true for P. pullata in the series HO, LD, HD. In LO more specimens of P. pullata were caught than could be expected on the basis of the correlation, mentioned above. More information may become available when the respective categories of spiders are analysed. Table 8 presents the mean numbers per category of both species caught in the four types of vegetation, per pitfall trap. Table 7. The mean numbers of both species per trap in the different types of vegetation Type Species prativaga pullata Total % prativaga of Total LO 4.2 7.0 11.2 40 LD 9.2 5.9 15.1 60 HO 7.6 4.3 11.9 61 HD LIZ 72 18.9 62 Although only a small number of females was caught, for both species the numbers, in percentages of the total, were slightly lower in type LO than in the others. In addition, only 6 % of the females of P. pullata caught in type LO carried an egg sac, whereas this percentage was 50—60 % in the other vegetation types. In P. prativaga 23—33 % of the females were with egg sac in all the vegetation types. Thus the higher numbers of P. pullata in vegetation type LO are only caused by the higher numbers of males and females without egg sac. In P. prativaga the relative proportions of the three categories of spiders is the same in the different types of vegetation, their numbers increasing from LO — HO — LD J. DEN HOLLANDER & H. Lor: Use of habitat by Pardosa 213 Table 8. The mean numbers of both species (per category) per trap in the different types of vegetation (Pc stands for ® with egg sac) Type Species prativaga pullata category a % Sof 2 Lc % Qcof a % & of Q Qc % Lcof total total 9 total total 9 LO 3.6 87 0.4 0.2 29 5.7 82 bia 0.1 6 HO 6.5 85 0.8 0.4 30 3:2 74 0.5 0.6 58 LD 1-3 81 1.4 0.4 23 4.3 73 0.8 0.9 53 HD 9.5 81 155 0.7 33 3:3, 74 0.9 0.9 50 — HD. Concerning males and females without an egg sac, there are differences between P. pullata and P. prativaga, the numbers of the former increasing from HO — LD — HD — LO. Females of P. pullata carrying an egg sac hardly occur in vegetation type LO. Thus, in P. prativara the largest numbers of the three categories of spiders occur in vegetation type HD, i.e. this vegetation type is prefered by males as well as by females, both with and without an egg sac. The same holds true for females of P. pullata with egg sacs. However, in this species males and females without an egg sac prefer vegetation type LO. CONCLUSIONS Of the numbers of both P. prativaga and P. pullata, taken by pitfall traps 80% are males. In P. prativaga 30 % of the females carry an egg sac, in P. pullata about 40 %. The numbers of P. prativaga amount to about 60 % of the total numbers caught. How- ever, in the samples taken by hand, males account for only 20 % of the numbers caught, in P. prativaga as well as in P. pullata. 60 % of the females of P. prativaga and 80 % of the females of P. pullata carry an egg sac. In this sample only 30 % of the spiders belong to P. prativaga. The difference in composition between the samples taken by pitfall traps and those taken by hand sampling reflects that females carrying an egg sac, as compared with both females without egg sac and males, show the lowest locomotory activity and occur more upon the vegetation. The different percentages of P. prativaga in the different samples are mainly caused by the higher activity of P. prativaga, as compared with P. pullata. Both P. prativaga and P. pullata occur throughout the investigated area. Towards the northeastern part of the field the numbers of both species increase, in P. pullata to a larger extent than in P. prativaga. Four different vegetation types, distinguished on the basis of their structure, occur in the study area. They are defined by the height and species of the composing plants, as well as by the percentages cover of both the low growing and high growing herb layers. As a consequence, the microclimatological conditions differ between the various vege- tation types (LO, LD, HO, HD). Both P. prativaga and P. pullata occur in all types of vegetation. In P. prativaga the numbers of males and females without an egg sac increase gradually from LO — HO — LD — HD; in P. pullata, on the contrary, from HO LD HD LO. In both 214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 4, 1972 species the numbers of females carrying an egg sac increase from LO — HO — LD — HD. Thus the prefered vegetation type for males and females of P. prativaga, as well as for females of P. pullata carrying an egg sac, is the high, densely structured vegetation (HD), and for males and females without an egg sac of P. pullata, the low, open structured type (LO). Discussion Former studies (Den Hollander, 1970, 1971) showed that in mixed populations interbreeding between P. prativaga and P. pullata might occur in some years. In those particular years, characterized by a cold spring, the seasonal barriers between both species are broken down. Laboratory studies showed that males of P. prativaga mate with females of P. pullata which results in living offspring. On the other hand, males of P. pullata do not mate with females of P. prativaga (cf. Den Hollander, 1972). Thus, interbreeding between the species studied under natural conditions can occur in only one direction, at least when both categories of spiders meet, i.e. when they are not isolated by a differential use of the habitat in mixed populations. The results of the present study show that both males and females of the species studied occur throughout the investigated area. Moreover, the numbers of P. prativaga caught per pitfall trap are positively correlated with those of P. pullata, and the dif- ferences between samples taken by hand and those taken by pitfall traps are similar in both species when the higher locomotory activity of P. prativaga males, as compared with males of P. pullata, is taken into account. Therefore, it may be concluded that males of the one species are not completely isolated from females of the other. Nevertheless, some differences in habitat preference between both males, and females without an egg sac (i.e. receptive females) of P. prativaga and P. pullata could be established. The largest numbers of P. pullata were obtained in vegetations, poorly grown with low growing plants. P. prativaga, on the other hand, was caught in the largest quantities in vegetations, densely grown with high growing plants. Besides, these two types of vegetations clearly differ concerning the values for temperature and relative humidity of the microclimatic conditions, as compared with those of the macroclimate. In the first type of vegetation temperature at the ground level is higher than that in the second type, mainly by the more direct radiation of the sun in the former. The opposite is true for the relative humidity. On the other hand, the influence of wind velocity and clouding on the microclimate will be less in the densely grown vegetations as compared with the other types. Probably, these results may be understood as a higher resistance to loss of water in P. pullata, as compared with P. prativaga. Indeed, unpublished observations in our laboratory showed that P. pullata survives at low humidities and high temperatures, for a much longer period than P. prativaga. The results concerning the preferent types of habitat for both species agree with the description of the characteristics of the habitats of P. pullata and P. prativaga given by previous authors (Dahl, 1908; Palmgren, 1939; Tambs-Lyche, 1940; Norgaard, 1945, 1951; Holm, 1947; Locket & Millidge, 1951; Wiebes, 1959; Tongiorgi, 1966; Vlijm & Kessler-Geschiere, 1967; Richter, Den Hollander & Vlijm, 1971; Den Hollander, 1971). According to these authors both species occur in moist fields overgrown to various extents. Especially Norgaard (1945, 1951) mentions that P. pullata prefers localities J. DEN HOLLANDER & H. Lor: Use of habitat by Pardosa 215 which are exposed both to direct radiation by the sun and to the influence of wind velocity. Several authors mention that P. pullata occurs in localities overgrown with low plants, whereas P. prativaga would occur in localities overgrown with high plants. In agreement with this, Richter et al. (1971) states that P. pullata is more adapted, measured in quantities of silk production, to open ground type vegetations and P. prativaga more to field type vegetations (see also introduction). These differences between P. prativaga and P. pullata mostly are gradual. In comparing distinct populations, both species occur in wide, overlapping ranges of vegetation types. Nevertheless, in pure as well as in mixed populations, the two species show a tendency to diverge to some extent in their use of habitat. This tendency to diverge may be caused by interspecific competition between both, closely related, species. This tendency may be inhibited, however, by the occurrence of interbreeding, as complete isolation mecha- nisms do not seem to exist between the species studied. REFERENCES Dahl, F., 1908. — Die Lycosiden oder Wolfspinnen Deutschlands und ihre Stellung im Haushalte der Natur. — Nova Acta Acad. Caes. Leop.-Carol. 88: 177—558. Hollander, J. den, 1970. — Morphological variation in Pardosa prativaga L. Koch, 1870, P. prativaga var. fulvipes Collett, 1875, and P. pullata Clerck, 1757 (Araneae, Lycosidae). — Tijdschr. Entomol. 113: 273—288. —, 1971. — Life histories of species of the Pardosa pullata group, a study of ten populations in the Netherlands (Araneae, Lycosidae). — Tijdschr. Entomol. 114: 255—281. , 1972. — Hybridization experiments in the Pardosa pullata group (Araneae, Lycosidae). — In preparation. Holm, A., 1947. — Svenske Spindelfauna. — Ent. Foreningen (Stockholm) 3: 1—48. Holm, A. & T. Kronestedt, 1970. — A taxonomic study of the wolf spiders of the Pardosa pul- lata group (Araneae, Lycosidae). — Acta ent. bohemoslov. 67: 408—428. Locket, G. H. & A. F. Millidge, 1951. — British spiders, i. — London. Norgaard, E., 1945. — Okologiske Undersogelser over nogle Danske Jagtedderkopper. — Flora Fauna, Silkeborg 51: 1—38. , 1951. — On the ecology of two lycosid spiders (Pirata piraticus and Lycosa pullata) from a Danish Sphagnum bog. — Oikos 3: 1—21. Palmgren, P., 1939. — Die Spinnenfauna Finlands. 1. Lycosidae. — Acta Zool. Fennica 25: 40—85. Richter, C. J. J., 1970. — Relation between habitat structure and development of the glandulae ampullaceae in eight wolfspider species (Pardosa, Araneae, Lycosidae). — Oecologia (Berl.) 5: 185—199. Richter, C. J. J., J. den Hollander & L. Vlijm, 1971. — Differences in breeding and mobility between Pardosa pullata (Clerck) and Pardosa prativaga (L. Koch) (Lycosidae, Araneae) in relation to habitat. — Oecologia (Berl.) 6: 318—327. Tambs-Lyche, H., 1940. — Die Norwegischen Spinnen der Gattung Pardosa Koch. — Avh. Norske Vidensk.-Akad. 1 (6): 1—59. Tongiorgi, P., 1966. — Italian Wolfspiders of the genus Pardosa (Araneae, Lycosidae). — Bull. Mus. Comp. Zool. 134: 275—334. Vliim, L. & A. M. Kessler-Geschiere, 1967. — The phenology and habitat of Perdosa monticola, P. nigriceps and P. pullata (Araneae, Lycosidae). — Journ. Anim. Ecol. 36: 31—56. Wiebes, J. T., 1959. — The Lycosidae and Pisauridae (Araneae) of the Netherlands. — Zool. Verh. 42: 1—78. taht ei very haat we » rape dpi rara ed zel ego ba grad vibe ht saint Loris “uit ol ike AHA apr à pme ed al nil dal atten, DOUTE rate Group lavica arts nb vai almeno GE 127° 4 TR PTL, ILES A ba Lie de are hi SA uber ines imp lie rit : | CHEZ onl ni Kol a learn) HE tti er BTN erh En u Da A iene conga ith Rp Ak AE CCL OC mee Le ent fired venata qu om anita he ei ar sla, ao app (osti Herma = i As » CT wy Nad nord gat els teal dog wenn wet ka vaderen aen TL SO 2 LT han boat dut rar mal (vial piro Oi enge m J uam Sue ae la Mo nee ju oa bisi ra Pe it etal: Lane lade meg | i i Ya |, 19 oa" Per + ï î brandde \ à AT a MT "nl ae ET” Wien eel iti id ial ACh ay À \ ot m US "il it os bebe we wy woe mb gen Wy) Ach bemerkt ged Nut deer er Oerle et “Titel ae ee (Ie een KALS, en Bi ART AMAR NES “la - iy ge ee TEL Re o M hae Al mill, VET bo tre MM BIJ Vue | Drei fn ar 4 er ni part ia? ally, WE veh ei CA nop | TE IT cae N werkd dS . Te i i | Jade val aarp, FLE i] i | (inl batt ade, Pup, Dlt Hi De re, OT le DADA TT PRE 1 (Vitti ce lor AE DUNK 18 nd Fine A pres "sa ws PNR a | ET Pur ul EN ro here adel Le „met: | è A The putin) sla LH | eg Mit Ta (hi ee eee | | dit) | gi Mane | inal a ; MUR bribi el hs ait i ru NY | ve Vat ge: a? t are’ Wad B yore ami à vue nt bl i i alent a Ue Wight oe! rel A | N ten § bel ki j h ly Mal Inte nal PT QE whee bts te tas il JM OT TITLE Se fin atid Wei alle OM Who TRA sl A Tal OE pit) in ve ee VUE i il Beer}. ballare i ler aa ‘ UT TO auf mir "No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.FI. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff, 1968 — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.FI. 70.—. No. 4. F. 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FURTHER STUDIES ON SOUTH AMERICAN GOMPHIDAE (ODONATA) by JABELLE Velp ABSTRACT The writer reports on new Gomphid material from South America. Among the 27 species recorded, Aphylla boliviana sp. n. is represented by females only, while of Phyllocycla vesta sp. n. and Phyl- locycla propinqua sp. n. both sexes are present. The latter was previously considered no more than an eastern form of Phyllocycla viridipleuris (Calvert). More individuals of other species, hitherto only known from the types, are recorded. The females of others are described for the first time. The genera Agriogomphus Selys and Cyanogomphus Selys are briefly re-diagnosed with partly new characters. CONTENTS Archaeogomphus nanus Needham — &, Surinam . . Agriogomphus jessei (Williamson) — & ®, Venezuela . Cyanogomphus conchinus Williamson — &, Surinam . Cyanogomphus uncatus Fraser — 2, Brazil . Epigomphus llama Calvert — & 2, Bolivia ‘ Desmogomphus tigrivensis Williamson — 9, TA | Apbylla boliviana spec. nov. — ®, Bolivia . Aphylla albinensis Belle — ®, French Guyana . Aphylla dentata Selys — 62, Surinam . Aphylla molossus Selys — 49, Brazil . ; 4 Aphylla distinguenda (Campion) — 88, ni : Aphylla theodorina (Navas) — 4, Brazil . 5 Phyllocycla modesta Belle — 9, Surinam . . Phyllocycla anduzei (Needham) — 49, Veneti Bod Boi : Phyllocycla titschacki (Schmidt) — 4, Bolivia . Na. Phyllocycla vesta spec. nov. — 4 9, Argentina . Phyllocycla argentina (Hagen in Selys) — 4 9, Argentina and Brazil è Phyllocycla viridipleuris (Calvert) — 4 9, Paraguay . Phyllocycla propingua spec. nov. — & Q, Brazil . Phyllocycla pallida Belle — &, Uruguay . i; ; Negomphoides undulatus (Needham) — 3, Brazil J Negomphoides lieftincki Belle — & 2, Bolivia . Zonophora batesi Selys — &, Brazil and Surinam . ; Zonophora calippus Selys — & ®, Brazil, Venezuela and Surinam : Zonophora spectabilis Campion — 2, Bolivia . Zonophora klugi Schmidt — 2, Brazil . : Zonophora surinamensis Needham — &, Surinam . References 217 page 218 218 221 221 221 221 222 224 224 225 226 226 227 227 227 227 230 233 235 236 236 236 236 237 237 238 238 239 218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 Since the publication of my studies on the South American Gomphidae in 1970, new material from this area was submitted to me for identification. The major part was sent to me by Prof. Dr. Minter J. Westfall, Jr., of the Univtrsity of Florida. The remaining material came from Dr. D. C. Geijskes, of the Leiden Museum, Dr. Kurt Günther, of the Berlin Museum, Dr. Borge Petersen, of the Copenhagen Museum and Dr. Douglas St. Quentin, of the Vienna Museum. I wish to express my gratitude to the colleagues mentioned. In the following pages records, descriptions and figures of this new material are presented. The names of the institutions in which the studied material is preserved are ab- breviated as follows. FSC — Florida State Collection of Arthropods, Gainesville. MZ — Museum of Zoology, University of Michigan, Ann Arbor. MC — Universitetets Zoologiske Museum, Copenhagen. ML — Rijksmuseum van Natuurlijke Historie, Leiden. MNB — Museum für Naturkunde der Humboldt-Universität, Berlin MNHW — National Museum of Natural History, Washington, D.C. NMW — Naturhistorisches Museum, Vienna. Archaeogomphus nanus Needham, 1944 Material. — Surinam: Nickerie River, Blanche Marie-val, 12.11.1971, 1 9, D. C. Geijskes (ML). Agriogomphus jessei (Williamson, 1918) Fig. 1—4 Material. — Venezuela: State of Tächira, La Fria, 15.1V.1920, 1 8, 1 9, J. H. and E. B. Williamson and W. H. Ditzler (FSC). There are some colour differences between the males from Tächira, Venezuela, and the male holotype from Cristalina, Colombia: 1. Rear of head yellowish; in holotype pale grey. 2. Dark antehumeral stripes on dorsum of pterothorax complete; in holotype inter- rupted in middle (this difference in the colour design is most striking). 3. First pale antehumeral stripe dull blue, other pale stripes of pterothorax green; in holotype pale markings on dorsum of pterothorax and mesepimeron dull blue. 4. Femora dark brown above, for the apical four-fifths on the first, for the apical half on the second, and for the apical one-third on the third; in holotype for the full length on the first, for the apical three-fourths or more on the second, and for the apical half or less on the third. 5. Pale markings of abdomen green; in holotype dull blue or greenish blue. 6. Anterior hamule brown, the edges black; in holotype anterior hamule entirely black. 7. Posterior hamule greenish, the extreme tip black; in holotype greenish with a constricted cross black bar at about one-third its length, the apical fourth black, darkest at apex fading basally. 8. Superior caudal appendages entirely yellowish; in holotype blackish basally fading to yellowish. 219 J. BELLE: South American Gomphidae Fig. 1—4. Agriogomphus jessei (Williamson), ®. 1, rear of head, dorsal part, with submedian pair of occipital spines; 2, vulvar scale and 9th sternum, ventral view; 3, diagram of pterothorax; 4, apical segments of abdomen, left lateral view. Fig. 5, 6. Cyanogomphus uncatus Fraser. 5, vulvar scale, 9 of 3.1.1941; 6, the same, ® of 7.1.1941. Fig. 7, 8. Desmogomphus tigrivensis Williamson, 2.7, apical segments of abdomen, ventral view; 8, the same, left lateral view 220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 Some venational features in the wings of the male from Tachira: Pterostigma covering 31/,—51/, cells. Antenodal and postnodal cross-veins of first series 7: 12—12:8/8: 10—10 : 7, in front and hind wings, respectively. Second primary antenodal cross-vein the fifth in each wing. Trigonal interspaces with two rows of cells. All supratriangles open. Four spines or small plates on posterior border of each hind wing basally to anal angle. The measurements of the male from Tachira: Total length 40 mm; abdomen 31 mm (incl. caud. app.); hind wing 23 mm; costal edge of pterostigma in front wing 2.8 mm. Female. — Total length 40.5 mm; abdomen 31.5 mm; hind wing 25.5 mm; costal edge of pterostigma in front wing 3 mm. Stouter than male, especially abdomen. Coloration most resembling the corresponding male from Tächira but vertex paler, pale middorsal stripe on hind lobe of prothorax broader and first pale antehumeral stripe green. Ninth abdominal segment one and a half times as long middorsally as midventrally (as is the case in the male). Sterna of segments 3 to 9 (except base of third sternum) blackish. Vulvar scale blackish, about two-fifths the length of ninth sternum, bifid in its apical one-third, the two divisions separated by an interval of 90°, their tips subacute, the bottom of the excision round. Caudal appendages pale yellow becoming darker to base, their surface minutely tuberculate, each tubercle with a pale hair. Lamina supra-analis three-fourths the length of caudal appendages. Wings larger than in male. Pterostigma covering 41/, cells. Antenodal and postnodal cross-veins of first series 8:11—12:8/8:10—9:8 in front and hind wings, respectively. Second primary antenodal cross-vein the fourth in left front wing and right hind wing, the fifth in other wings. Trigonal interspaces with two rows of cells. All supratriangles open. A single row of cells behind Cu2 in front wing, the same area in hind wing with two rows of cells. Hind wing with four paranal cells and two postanal cells, the fourth paranal cell is the first postanal cell. Remark: The genus Agriogomphus is nearest allied to Cyanogomphus from which it differs in the following particulars: 1. Occiput with a pair of submedian spines near foramen; no occipital spines in Cyanogomphus. 2. Apical half of posterior genital hamule of male slender, tapering, and bare; stout, more or less sickle-shaped, and tufted with hairs in Cyanogomphus. 3. Inferior caudal appendage of male slender, the two branches short, reaching to a point just beyond spurs of superior appendages, and armed with a minute, subapical superior tooth; in Cyanogomphus very stout, with long strong branches, reaching to a point far beyond spurs of superior appendages, and not armed with a subapical superior tooth. 4. Male anal triangle in hind wing absent (ill-defined); well-developed and three- celled in Cyanogomphus. Calvert's twmens also possesses a pair of occipital spines and for that reason that species can be referred to Agriogomphus. The occipital spines are better developed in the females than in the corresponding males; they are short in A. sylvicola and A. ericae, well-developed and conspicuous in A. tumens and A. jessei. J. BELLE: South American Gomphidae 221 Cyanogomphus conchinus Williamson, 1916 Material. — Surinam: Nickerie River, Stondansi, 18.11.1971, 1 g, D. C. Geijskes (ML). A teneral male, partly crushed. Cyanogomphus uncatus Fraser, 1947 Fig-25 06 Material. — Brazil: Santa Catarina, Nova Teutonia, 3.1.1941, 1 9; 7.1.1941, 1 © , Fritz Plaumann (MZ). The tip of the vulvar scale in the female dated 3.1.1941, is narrowly excised in middle for nearly half the length of the vulvar scale, that in the female dated 7.1.1941, is cleft for only two-fifths of the length of the vulvar scale. Further the frons in these two females is slightly broader than that of the female from Rio Grande do Sul, recorded in my 1970 paper on the South American Gomphidae. Finally, the female from Rio Grande do Sul possesses a small but distinct tubercle below each of the swollen superior portions of the occiput. These occipital tubercles are low in the female from Santa Catarina, collected on 7.1.1941, and practically wanting in the female, collected on 3.1.1941. Epigomphus llama Calvert, 1903 Material. — Bolivia: Dept. Cochabamba, Prov. Chapare, El Palmar (1600 m), 1 9, R. Steinbach; Cristal Mayu (600 m), 1 & (FSC). The female is teneral and in bad condition, crushed over the entire length of the body. Desmogomphus tigrivensis Williamson, 1920 Eion 78 Material. — Guyana: Kaieteur Ravin, 11.1V.1912, 1 9, J. M. Geddes (MZ). The present female is somewhat larger than the corresponding male described from the same country by Williamson, but otherwise agreeing in all respects other than those of sex. The vulvar scale is distinctly shorter than in the female of Desmogomphus paucinervis (Selys) and this feature forms the evidence for the specific difference of these two species. Female. — Total length 45 mm; abdomen 34 mm; hind wing 29 mm; distance on hind wing from nodus to pterostigma 11.5 mm; width of hind wing at level of arculus 8.8 mm; costal edge of pterostigma in front wing 3.1 mm, in hind wing 3.5 mm. Face predominantly brown but base of mandibles externally, genae, most of labrum (except for free border and base) and lateral sides of postclypeus greenish. Superior surface of frons greenish but dark brown in middle and along base. Vertex dark brown, behind lateral ocelli somewhat bulbous and greenish. Occipital plate brown, low, the breadth about six times the middorsal length, the posterior margin fringed with long, dark brown hairs. Rear of head brownish, becoming yellowish below. Labium and ad- jacent mouth parts pale yellowish. Prothorax brown. Pterothorax brown with green stripes, its pattern similar to that of male except for third lateral brown stripe which consists of two isolated brown stripes separated by the metapleural suture. Legs slender. Femora brownish but first femora darker on dorsal side, and middle 222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 and posterior femora becoming darker towards knees. Tibiae, tarsi and claws dark brown. Abdomen predominantly dark brown, with lighter spots on sides of segments 8 and 9. Pale colours greenish or yellowish. Segment 1 laterally, middorsum of segment 2 and apical half of middorsum of segment 3 pale. A pale middorsal line on segments 4 to 7, becoming broader at base of segment 5 to 7, pale baso-lateral spots on segments 4 to 7, followed by a pale lateral spot on segments 6 and 7. Anal appendages yellow. Vulvar scale and sterna of segments 8 and 9 brown. Tip of vulvar scale deeply excised for two-fifths the length of scale, the divisions reaching to a point about halfway along ninth sternum. Wings slightly brownish. Venation brown, including frontal margin of costa. Pterostigma light brown, surmounting 3—4 cells. Basal subcostal cross-vein wanting. Antenodal and postnodal cross-veins of first series 13 : 14—15 : 11/11 : 10—10 : 10 in front and hind wings, respectively. Second primary antenodal cross-vein, the sixth in left front wing, the fifth in other wings. Intermedian cross-veins 5—5/3—3 in front and hind wings, respectively. All supratriangles, subtriangles and triangles open except for triangle in hind wings which is two-celled. Front wing with a single row of cells in anal field proximal to level of triangle and in area posterior to Cu2. Trigonal inter- space in front wings starting with two rows of cells from triangle out, seven cells long, in anterior row. Trigonal interspace in hind wings starting with a row of 3 (left) and 4 (right) cells against triangle, followed by two rows of cells, four cells long, in anterior row. Each hind wing with 5 paranal cells and 3 postanal cells, the fifth paranal cell is the first postanal cell. Three rows of cells behind Cu2 in hind wing. Aphylla boliviana spec. nov. Fig. 9, 10 Material. — Bolivia: Dept. Santa Cruz, Prov. Ichilo, Rio Dolores, 3 km east of Buena Vista, 23.111.1960, 1 9, R. B. Cumming, Quiroga, and R. Rodriguez (holo- type); Dept. Cochambamba, Prov. Chapare, Palmar, 30 km from Villa Tunari on road toward Cochabamba, collections at a lake, altitude 1935 ft, 24.X.1959, 1 9; Dept. Santa Cruz, Prov. Ichilo, pond stream locality, 18 km west of Buena Vista, 8 km west of San Carlos, 16.1II.1960, 1 9; 17.111.1960, 2 Q, all R. B. Cumming (FSC). The nearest relative of this species is, perhaps, Aphylla producta Selys. The female agrees by the completely unexpanded ventro-tergal margins of the abdominal segments 8 and 9, but it differs from all other members of the genus by the conspicuous orange band along the lateral inferior borders of the ninth abdominal segment. Female (holotype). — Total length 58.5 mm; abdomen 44 mm; hind wing 35 mm; costal edge of pterostigma in front wing 4.5 mm. Face dark brown, the following green: Base of mandibles externally, a symmetrical pair of oblong spots on labrum, anteclypeus, concave areas and lateral posterior portions of postclypeus. Anterior margin of labrum orange and fringed with orange bristles. Superior surface of frons dark brown at base, green anteriorly, the green band at level of antennae nearly three times as wide as in middle and twice as wide as basal band. Vertex dark brown, greenish on concave area behind postocellar ridges. Occipital plate ‘greenish, its hind margin brown and fringed with brown hairs which are in length about half the middorsal width of the occipital plate. Rear of head dark brown above, greenish on temporae. Labium and adjacent mouth parts greenish. J. BELLE: South American Gomphidae 223 15 18 Fig. 9, 10. Aphylla boliviana spec. nov., 9 holotype. 9, diagram of pterothorax; 10, vulvar scale. Fig. 11, 12. Aphylla molossus Selys, 9. 11, diagram of pterothorax; 12, vulvar scale. Fig. 13, 14. Aphylla distinguenda (Campion), ®. 13, diagram of pterothorax, Posadas; 14, vulvar scale, San Fernando. Fig. 15. Aphylla dentata Selys. Vulvar scale of allotype. Fig. 16. Aphylla theodorina (Navás), 4. Diagram of pterothorax, Minas Gerais. Fig. 17, 18. Phyllocycla anduzei (Needham). 17, vulvar scale, © of 12.III.1960; 18, the same, ® of 23.III.1960 Prothorax brown, middle lobe with a green spot on dorsum and a green spot on each side. Pterothorax dark brown with green stripes; its markings shaped as shown in Fig. 9. Middorsal carina brown, posterior part of metepimeron and metapostepimeron less obscure than other dark stripes of pterothorax. Posterior margin of metaparapteron more acute than is normally the case in Aphylla (in Phyllocycla more or less undulate). Femora reddish brown, the inner side of anterior pair green. Tibiae, tarsi and claws biackish. Posterior tarsi two-thirds the length of posterior tibia. Abdomen predominantly reddish brown. Sides of ninth segment with an orange band along lateral borders, this band regularly widening to base of segment. Tenth segment 224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 orange but swollen basal one-sixth portion brown and dorso-apical rim blackish. Dorso- apical rim with a conspicuous pair of paler (polished) spots on middorsum. Width of dorso-apical rim about one-seventh the length of tenth segment. Dorsal posterior margin of tenth segment denticulated, the denticles smaller in middle. Appendages nearly as long as tenth segment, the extreme tips outcurved. Vulvar scale short, about one-sixth the length of ninth sternum, its posterior margin deeply and widely excised, V-shaped. The relative length of the three apical segments of abdomen is about 18 : 13 : 10, with anal appendages 9 at the same scale. Wings hyaline. Venation dark brown, including frontal margin of costa. Pterostigma yellowish brown, covering 6—8 cells. Basal subcostal cross-vein present. Antenodal and postnodal cross-veins of first series 11 : 20—21 : 11/11 : 15—13 : 12 in front and hind wings, respectively. Second primary antenodal cross-vein the seventh in front wings, the sixth in left hind wing, and the fifth in right hind wing. Intermedian cross-veins 11—11/6—6 in front and hind wings, respectively. Trigonal interspace starting with a row of three cells against triangle followed by two rows of cells, in front wings 6 (right) and 8 (left) cells long, in anterior row, in hind wings 4 cells long, in anterior row. Supratriangles two-celled. Triangle in front wings three-celled. Subtriangle in front wings and triangle in hind wings two-celled. Subtriangle in hind wings open. Four rows of cells behind Cu2 in front wings, four to five rows of cells behind Cu2 in hind wings. Anal loop in hind wings two-celled. Hind wings with 4 paranal cells and 4 postanal cells, with the veins A3 and A2 parallel, and with the second anal inter- space starting with a single large cell against anal vein. Male unknown. There is a striking difference in size between the five females recorded here; the holotype is the smallest of the series (but the best preserved specimen); the larger specimens are much more robust. The dimensions of the largest female are: total length 66.5 mm; abdomen 50 mm; hind wing 41.5 mm; costal edge of pterostigma in front wing 5.5 mm. The form of the vulvar scale is variable, from deeply and widely V-shaped to nearly semicircular. In two females the pale frontal band of the superior surface of the frons is much broader than in the others. Some venational features in the wings of the four paratypes are as follows. Front wings: four supratriangles three-celled, other two-celled; all triangles three-celled; two subtriangles three-celled, others two-celled. Hind wings: all supratriangles two-celled; one triangle three-celled, others two-celled; all subtriangles open; in two hind wings the second anal interspace starts with a single cell against the anal vein, in others with two cells. Aphylla albinensis Belle, 1970 Material. — French Guiana: Saint Laurent, 1 9, A. Kricheldorff (MNB). A teneral specimen with the abdomen crushed over its entire length. Saint Laurent is a town at the Marowijne River, opposite Albina, from where the species was described. Aphylla dentata Selys, 1859 Material. — Surinam: 2 g, 1 9 (MNB). The female lacks the sixth abdominal segment. J. BELLE: South American Gomphidae 225 Aphylla molossus Selys, 1869 Fig. 11, 12 Material. — Brazil: Rio Purus, Arima, 22.X1.19??, 1 g, 1 9, S. M. Klages (FSC). The female of this species has not been described. The present female has the subtriangle in front wing three-celled (in male recorded here, two-celled), that in right hind wing two-celled, and that in left hind wing possesses a partly developed cross- vein. Female. — Total length 59 mm; abdomen 45 mm; hind wing 35.5 mm; costal edge of pterostigma in front wing 4.5 mm. Face brown but genae, base of mandibles externally and anteclypeus green, labrum with a symmetrical pair of green oblong spots, and postclypeus with a green posterior spot on each side. Superior surface of frons brown at base, green anteriorly. Vertex blackish brown, reddish brown behind lateral ocelli. Occipital plate greenish, its posterior margin straight and fringed with short hairs. Rear of head brown, becoming greenish below. Labium and adjacent mouth parts pale greenish. Prothorax brown, its hind lobe with green central spot and a green spot on each side. Pterothorax brown with green stripes. Middorsal carina green. Green anterior meso- thoracic “half collar” interrupted in middle. First green antehumeral stripe not con- nected with green mesothoracic “halr collar and with well-developed second green antehumeral stripe immediately in front of humeral suture. A well-developed green stripe on mesepimeron, on metepisternum and on metepimeron. Femora reddish brown. Tibiae, tarsi and claws blackish. Abdomen dark brown. Dorso-apical rim of tenth segment paler, its middorsal width about one-seventh the length of segment 10. Lateral expansions of segments 8 and 9 narrow, equal in width and denticulated. Vulvar scale about one-sixth the length of ninth sternum, its posterior median excision widely V-shaped with a round bottom. Wings slightly brownish. Venation dark brown including frontal margin of costa. Pterostigma light brown, surmounting 7—91/ cells. Basal subcostal cross-vein present. Antenodal and postnodal cross-veins of first series 17 : 25—23 : 17/19 : 18—20:19 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth in left hind wing, the eighth in left front wing, and the seventh in right wings. Inter- median cross-veins 13—14/8—8 in front and hind wings, respectively. Supratriangle in left front wing three-celled, in other wings two-celled. Triangle and subtriangle in front wings three-celled. Triangle in left hind wing three-celled, in right hind wing two-celled. Subtriangle in left hind wing open but with an undeveloped cross-vein, in right hind wing two-celled. Trigonal interspace in front wings starting with a row of three cells against triangle followed by three rows of cells, here and there inter- rupted by two cells, one cell enlarged. Trigonal interspace in hind wings starting with a row of three cells against triangle followed by two cells (right), or a row of two cells (left), two cells long. Five paranal cells and five postanal cells in each hind wing. Five rows of cells behind Cu2 in hind wings. Remark: The pale spots on the labrum and the postero-lateral pale spots on the postclypeus are weakly developed in the male. In the holotype male the spots on the labrum are obviously obliterated owing to postmortem changes. 226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 Aphylla distinguenda (Campion, 1920) Fig. 13, 14 Material. — Argentina: Posadas, 6.111.1909, 1 9 (MC); Prov. Buenos Aires, Dique Lujan, 31.X11.1957, 1 g, 1 9, Hellmut G. Gloger (FSC); San Fernando, I, II, 111.1967, 1 8, 2 9, J. Daguerre; Tigre, 17.11.1968, 2 g', O. S. Flint, Jr. (MNHW). The female of this species has been described under Aphylla dentata by Ris in 1904 and a diagram of its pterothorax was published by him in 1913. The female described below is from the Copenhagen Museum. Female. — Total length 58 mm; abdomen 44.5 mm; hind wing 34.5 mm; costal edge of pterostigma in front wing 5 mm. Similar to male as regards stature and general coloration. Colour design of ptero- thorax slightly differing from that of holotype, male, and shaped as shown by Fig. 13. The brown on dorsum of pterothorax obscure but distinctly paler between first green antehumeral stripes which are vaguely connected below for inner half of width with green mesothoracic “half collar’. Middorsal carina green. Second green antehumeral stripe immediately in front of humeral suture weakly developed and represented only by two narrow green spots, one anterior (below) and one near mid-height. Ventro-tergal margin of abdominal segments 8 and 9 very narrowly expanded and denticulated at apex. Hind dorsal margin of segment 10 with denticles, those in middle smaller. Caudal appendages a trifle shorter than segment 10, conical with the tips slightly curved outward. Dorso-apical rim of tenth abdominal segment one-eighth the length of segment. Vulvar scale one-sixth the length of ninth sternum, its posterior margin deeply and widely excised, V-shaped. Frontal margin of costa yellow. Antenodal and postnodal cross-veins of first series 10 : 20—19 : 12/13 :13—14 : 14 in front and hind wings, respectively. Second primary antenodal cross-vein the seventh in front wings, the sixth in hind wings. Triangle in front wings three-celled, the dividing cross-veins tri-radiate from centre. Triangle in hind wings and subtriangle in front wings two-celled. Subtriangle in hind wings open. Anal loop in hind wings two-celled. Hind wings with five (right) and four (left) paranal cells. Four postanal cells in each hind wing. In the other specimens the second pale antehumeral stripe is also weakly developed and its upper portion is not or hardly perceivable. In the female from San Fernando the triangle in the right hind wing is open, the subtriangle in the left front wing is also open but possesses a partly developed cross- vein, the excision of the vulvar scale is perfectly semicircular, and the green markings are better developed than in the other specimens and this applies particularly to the sides of the pterothorax. Aphylla theodorina (Navas, 1933) Fig. 16 Material. — Brazil: Rio Grande do Sul, Pelotas, 2.11.1955, 1 &, C. Biezanko (FSC); Minas Gerais, XII, 1967, 1 g', V. da Cajoa and S. do Cipo (NMW). The two males have the dorsal posterior margin of the tenth abdominal segment armed with denticles at level of bases of superior appendages. The male from Rio J. BELLE: South American Gomphidae 227 Grande do Sul is teneral. The one of Minas Gerais has the pale colours greenish and the pterothorax with reduced pale markings, the second pale antehumeral stripe immediately in front of the humeral suture developed for its dorsal half only, the second pale lateral stripe (on the metepisternum) weakly developed, the metapostepimeron entirely dark brown, etc. Also in the male from Minas Gerais the posterior margin of the metaparapteron is nearly straight, in the specimens from Rio Grande do Sul pointed, as is normally the case in Aphylla. Phyllocycla modesta Belle, 1970 Material. — Surinam: Nickerie River, Lombokval, 8.11.1971, 1 9, D. C. Geijskes (ML). An old specimen, as clearly appears from the very brown wings. Phyllocycla anduzei (Needham, 1943) Fig. 17, 18 Material. — Venezuela: 2 8 (MNB);1 & (coll. of author), all F. Kummerow. — Bolivia: Dept. Santa Cruz, Prov. Ichilo, Rio Dolores, 3 km east'of Buena Vista, 12.111.1960, 1 ©, R. B. Cumming, Quiroga, and R. Rodriguez; 23.111.1960, 1 9, R. B. Cumming (FSC). The two females from Bolivia may belong to Phyllocycla anduzei recorded from that country by me in 1970. The vulvar scale of these two specimens differs from that of the allotype female from Venezuela. In the allotype the interval between the lobes of the posterior excision is 90°, in the two females recorded here, 60°. But between the two females from Bolivia differences exist also in the shape of the excision. It is V-shaped with a flat bottom in the female of March 12 and sharply V-shaped in the female of March 23. In the males from Venezuela and Bolivia differences exist also in the form of the ventro-tergal margin of the seventh abdominal segment. Phyllocycla titschacki (Schmidt, 1952) Material. — Bolivia: Dept. La Paz, Prov. Nor Yungas, 1 km north of Caranavi (rapid mountain stream), 13.IV.1960, 2 4, R. B. Cumming (coll. of author). Phyllocycla vesta spec. nov. Fig. 19—25 Material. — Argentina: Prov. Buenos Aires, San Fernando, I, II, 1II.1967, 27 3, 20 ®, J. Daguerre (holotype male, allotype female, and paratypes); Punta Chica, San Fernando, 15.11.1968, 2 g', 1 9, O. S. Flint, Jr. (paratypes, MNHW). Closely related to Phyllocycla gladiata (Hagen in Selys) from Pernambuco, Brazil, but larger and paler, with a quite different colour design of the pterothorax and ab- domen (the first pale antehumeral stripes broader, etc.). No other congeneric species has the exfoliations of the eight abdominal segment better developed, in the male they resemble greatly those of Ph. gladiata, but the apical portions are broader. Also the 228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 male genitalia on the second abdominal segment approach those of Ph. gladiata. The superior caudal appendages of the male are larger and stouter, and the subapical in- turned rim of the upper margin of these appendages is much better developed than in Ph. gladiata. The vestigial inferior caudal appendage of the male is relatively long, and its slender tip is distinctly excised, V-shaped. Male (holotype). — Total length 55 mm; abdomen 42 mm (incl. caud. app.); hind wing 30 mm; costal edge of pterostigma in front wing 4.4 mm. A greenish specimen marked with dark brown stripes and spots, but with the apical portion of the abdomen, including the lamellate expansions and caudal appendages, yellow. Face pale greenish but vertical part of frons somewhat obscure in middle and along postclypeus. Superior surface of frons largely pale greenish, brownish along base. Vertex brown, the concave area behind and between postocellar ridges yellow. Occipital plate yellow, its hind margin blackish brown, nearly straight and fringed with rather short pale hairs. Rear of head yellow behind occipital plate, dark brown on top behind compound eyes, brownish yellow on temporae. Labium and adjacent mouth parts pale yellow. Prothorax brownish, the sides and posterior dorsal part of middle lobe greenish. Pterothorax greenish with dark brown stripes; its pattern shaped as shown in Fig. 20. Middorsal carina greenish. Femora yellowish, becoming dark brown towards knees. Ventral (inner) sides of anterior and middle pair of femora greenish. Seven spines on distal half of antero- inferior row of posterior femur, the longest spines nearly one-third the diameter of femur. Tibiae, tarsi and claws blackish. Abdominal segments 1 and 2 greenish below, including auricles, dark brown above level of auricles, but middorsum of segments greenish again. Segments 3 to 6 and basal half of segment 7 greenish, apical half of segment 7 and segments 8, 9 and 10, including lamellate expansions and caudal appendages, yellow. A dark brown stripe on each side of apical half on segment 3, and from near the base to apex on segments 4, 5 and 6. Dorsum of segments 3 to 8 with a pair of dark brown spots near apex. A middorsal stripe on segments 3 to 7, isolated and confined to middle portion of segment 3, extending to apex of segment and touching the apical spots on 4 and 7, broadly confluent with these spots and forming with these a lance-shaped middorsal spot, from base to apex on 5 and 6. Segment 9 with a brown marking on each side. Segment 10 dark brown along dorsal posterior margin and on middorsum of base. Articulations between segments 3—4, 4—5, 5—6, 6—7 and nodulus between segments 7—8, 8—9, 9—10 dark brown. Dorso-apical rim of tenth abdominal segment about one- fourth the length of segment on middorsum, about one-third the length of segment on sides. Genitalia on second abdominal segment shaped as shown by the accompanying figures. Tip of posterior hamule distinctly stouter than in Ph. gladiata. Posterior ex- cavation of hood of penial peduncle with a distinct tooth on bottom. Wings clear, venation brown but frontal margin of costa yellow. Pterostigma yellowish brown, surmounting 5—6 cells. Basal subcostal cross-vein present. Antenodal and postnodal cross-veins of first series 9 :13—15 :9/11:11—12:9 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth in right front wing, the fifth in other wings. Intermedian cross-veins 9—9/6—6 in front and hind J. BELLE: South American Gomphidae 229 21 LL AWM Fig. 19—25. Phyllocycla vesta spec. nov., holotype and allotype. 19, 10th abdominal segment and caudal appendages of 4, dorsal view; 20, diagram of pterothorax, 4; 21, apical segments of ab- domen and caudal appendages of &, left lateral view; 22, genitalia on second abdominal segment, ventral view; 23, the same, right lateral view; 24, vulvar scale, ventral view; 25, apical segments of abdomen and caudal appendages of ©, left lateral view 230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 wings, respectively. All triangles, subtriangles and supratriangles two-celled except for subtriangle in hind wings, which is open. Trigonal interspace in front wings starting with a row of two (left) and three (right) cells against triangle followed by two rows of cells, 6 (left) and 8 (right) cells long, in anterior row. Trigonal interspace in hind wings starting with a row of three cells against triangle, followed by two rows of cells, 6 cells long, in anterior row. Hind wing: 4 paranal cells and 5 postanal cells; anal loop two-celled; distal portion of A2 convergent with A3; second anal interspace (between A2 and A3) with a single row of cells; 4 rows of cells behind Cu2; anal triangle four-celled. Female (allotype; a somewhat old specimen). — Total length 54 mm; abdomen 41 mm; hind wing 32 mm; costal edge of pterostigma in front wing 4.6 mm. Similar to male as regards stature and coloration. The slight colour differences confined to abdomen. There is a dark brown stripe on each side of basal half of segment 7 and the dorsum of segments 8 and 9 is obscure brown. Dorso-apical rim of tenth abdominal segment one-fifth the length of segment. Lamellate expansions of segment 8 well-developed and yellow. Wings brownish. Supratriangle in left front wing three-celled, in other wings two- celled. Subtriangle in front wings two-celled, in hind wings open. Triangle in front wings three-celled, in hind wings two-celled. Antenodal and postanal cross-veins of first series 10 : 16—17 : 9/11 :13—11:10 in front and hind wings, respectively. Second primary antenodal cross-vein the seventh in right front wing, the fifth in right hind wing and the sixth in left wings. Intermedian cross-veins 8—9/6—7 in front and hind wings, respectively. Second anal interspace in hind wings with two rows of cells. Five paranal cells and five postanal cells in hind wings. All specimens have the subtriangle in the hind wing open. All 29 males have the supratriangles, the triangles in the front wings and the anal loop in the hind wings two-celled, except one male which has the triangle and the anal loop in the right hind wing one-celled, but is seems that this wing was damaged during the larval stage. : The venational features of the 21 females were tabulated with the following results: supratriangle two-celled 95%, three-celled 5%; subtriangle in front wing open 2%, two-celled 98%; triangle in front wing two-celled 69%, three-celled 31%; triangle in hind wing two-celled 100%; anal loop in hind wing two-celled 91%, three-celled 2%, no anal loop in hind wing 7%. Phyllocycla argentina (Hagen in Selys, 1878) Fig. 26—29 Material. — Argentina: Cördoba, 1 9 (lectotype by present designation, MCZ No. 12382); Prov. Buenos Aires, San Fernando, I, II, III.1967, 2 8, 3 9, J. Daguerre; Punta Chica, San Fernando, 15.11.1968, 3 3, 1 9, OS: ÆFlint Alt Tigre, 17.11.1968, 1 9, O. S. Flint, Jr. (MNHW); Rosaria de Sta Fé, 1 &, Hubrich (MNB); Corrientes, 3.III.1909, 1 4, Esben-Peterson (MC). — Brazil: Porto Alegre, 1 9, Hensel (MNB). Described from a single pair from Cérdoba, Argentina, both individuals incomplete, lacking the apical segments of the abdomen. The holotype, a very teneral male, could not be located in the Hagen collection at Cambridge; it is probably lost. For that reason the female allotype is designated lectotype. The female type bears the pin labels J. BELLE: South American Gomphidae i] ISS] Fig. 26—29. Phyllocycla argentina (Hagen in Selys), 9. 26, diagram of pterothorax, Porto Alegre; 27, vulvar scale, Porto Alegre; 28, left posterior femur, left lateral view, Porto Alegre; 29, the same, lectotype. Fig. 30, 31. Negomphoides lieftincki Belle, &. 30, diagram of dorsum of ptero- thorax, Prov. Nor Yungas; 31, the same, Prov. Ichilo “Argentina”, “Cordova Argent.”, “Cyclophylla argentina Hag.”, “MCZ”, and “Type 12382”. It is redescribed below. Female (lectotype; abdominal segments 6—10 lost). — Abdominal segments one to five 21.5 mm; hind wing 33.5 mm; costal edge of pterostigma in front wing 5.2 mm. Face pale yellow. Superior surface of frons becoming brownish at extreme base. Scape yellow, pedicel brown, rest of antennae blackish. Vertex pale brownish-yellow. Oc- cipital plate yellow. Posterior margin of occiput nearly straight, with the slightest trace of a median excision, fringed with brown hairs which are about three-fourths as long as the middorsal width of occipital plate. Rear of head brownish-yellow, becoming somewhat more obscure above. Labium and adjacent mouth parts yellow. Prothorax yellow, the hind lobe slightly brownish. Pterothorax predominantly yellow, with brown markings. Dorsum brown with yellow stripes. Middorsal carina yellow. First pale antehumeral stripe rather broad, more or less parallel-sided, conjoined below with pale anterior mesothoracic “half collar’, the latter interrupted in middle. Second pale antehumeral stripe well-developed, not joined 232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 with first pale antehumeral stripe. There is a narrow yellow stripe covering the humeral suture. First brown lateral stripe narrow. Second brown lateral stripe weakly developed, interrupted near its upper end, which is represented by a spot. Third brown lateral stripe very weakly developed, being represented by a narrow metepimeral stripe only. Femora yellowish, anterior and middle pair of femora with a brown stripe on distal half of outer side. Spines on antero-inferior margin on distal side of posterior femur about one-third as long as diameter of femur. Tibiae dark brown with yellow dorsal side. Tarsi brown, middle joint yellow on dorsal side. Claws dark brown. Abdominal segments 1 to 5 predominantly yellow. Intersegmental articulations between segments 2 and 3 dark brown posteriorly, that between segments 3—4, 4—5, and 5—6 largely dark brown. Segments 1 to 3 yellow on middorsum for entire length of segment. Sides of segments 1 and 2 becoming brownish towards dorsum. Base of segments 4 and 5 yellow. Dorsum of segments 4 and 5 (except base) brown, the mid- dorsal brown stripe becoming wider to rear. Sides of segments 3 to 5 with transverse furrow and posterior submedian dorsal scar dark brown. Segments 4 to 5 with a long dark brown spot parallel to ventral margin. Middorsum of segments 3 to 5 armed with minute black denticles. Wings hyaline. Venation brown but frontal margin of costa yellow. Pterostigma yellow, covering 515—7 cells. Basal subcostal cross-vein present in right front wing, wanting in other wings. Antenodal and postnodal cross-veins of first series 9 : 16—18 : 11/11 :14—13 :11 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth. Intermedian cross-veins 10—10/7—6 in front and hind wings, respectively. Trigonal interspace of right hind wing with three rows of cells, three cells long from triangle out, followed by two rows of cells, three cells long. Trigonal inter- space in other wings starting with a row of three cells against triangle followed by two rows of cells. All triangles, subtriangles and supratriangles two-celled, except for sub- triangle in right hind wing, which is open. Five paranal cells and four postanal cells in each hind wing. In my 1970 paper, on the Navas types of Neotropical gomphines, it was stated that the specimens of Ph. argentina in the eastern parts of Argentina and the southern parts of Brazil are distinguished from the nominotypical form from Cordoba in having the tibiae and tarsi entirely black, and in having the brown lateral stripes of the pterothorax better developed, the second brown lateral stripe complete, the third brown lateral stripe represented by two isolated stripes (separated by the metapleural suture), and with an additional brown spot just above the mid-height of the metepimeron. Since I have been able to study the good series of this species from Buenos Aires, I can state that, with regard to the pattern of the pterothorax, the specimens of the eastern form are not uniformly coloured. In one specimen the second brown lateral stripe is very weakly developed, and in several other specimens the third brown lateral stripe is represented or practically represented by the narrow metepimeral stripe only. Of all specimens taken in Buenos Aires, however, the tibiae and tarsi are entirely black. Also, in the specimens from the type locality Cordoba, the pterostigma is longer. In the males recorded here the costal edge of the pterostigma in the front wing ranges from 3.9 to 4.4 mm, in the females recorded here (except the lectotype) from 4.2 to 4.6 mm. It is not improbable that, when the species is better known from Cérdoba, the eastern form will prove to be a subspecies of Ph. argentina. J. BELLE: South American Gomphidae 233 The lectotype female lacks the apical segments of the abdomen. In the other females recorded here some variability has been found in the denticulation of these segments. This denticulation is normally as follows: 1. Tenth abdominal segment without denticles in middle portion of dorsal posterior margin. 2. Ventro-tergal margin of eighth abdominal segment with denticles in apical half only. 3. Ventro-tergal margin of ninth abdominal segment without denticles. In the teneral female from Corrientes (Senckenberg Museum, no. 14484) the tenth abdominal segment is armed with denticles along the entire dorsal posterior margin, the ventro-tergal margin of the eighth abdominal segment has denticles on the apical three-fourths and that of the ninth abdominal segment possesses denticles on the basal submedian portion. The allotype, female, of Ph. eugentae (Navas) from Sierra de Cördoba, which is possibly conspecific with Ph. argentina, has the apical segments of the abdomen denticulated normally. Also the female from Porto Alegre has the apical segments of the abdomen normally denticulated. The subtriangle in the left hind wing of this female is two-celled and that in the right hind wing possesses a not wholly developed cross-vein. Phyllocycla viridipleuris (Calvert, 1909) Fig. 32—34 Material. — Paraguay: Dept. of Guaira, Villarrica, 15.XII.1938, 1 9; Paso- Yobay, 20.1.1939, 1 g (FSC). The male of this species has the apical inferior angles of the tenth abdominal segment folded under and applied against the sternum. In the present male the first pale antehumeral stripe is much broader than in the holotype and the upper end of this stripe is confluent with the second pale antehumeral stripe, immediately in front of the humeral suture. The female recorded here and hitherto unknown from the type locality, Paraguay, is described below. Female. — Total length 50 mm; abdomen 38.5 mm; hind wing 31.5 mm; costal edge of pterostigma in front wing 4.1 mm. Similar to the male holotype with regard to the stature and general coloration, but face paler. Bases of mandibles externally, genae and postclypeus pale yellow. Labrum, anteclypeus and frons pale greenish. Vertex and occipital plate brown. Prothorax entirely green. Pterothorax, as in holotype, largely yellowish green, with dark brown markings. Legs as in holotype with outer sides of tibiae yellowish. Abdomen predominantly dark brown but middorsum and sides of segments 1 and 2, and middorsum of base of segment 3, yellowish. Ventro-tergal margins of segment 8 denticulated on apical half, slightly expanded, being twice as wide as those of segment 9. Dorso-apical rim of tenth segment one-sixth the length of segment. Stylets nearly as long as tenth segment. Vulvar scale brown, about two-fifths the length of ninth sternum, its posterior margin widely and deeply excised for three-fourths the length of the vulvar scale. Wings slightly brownish-yellow at base. Venation brown but frontal margin of costa 234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 Fig. 32—34. Phyllocycla viridipleuris (Calvert). — 32, 10th abdominal segment and caudal appen- dages, &, ventral view; 33, vulvar scale; 34, apical segments of abdomen, 9, left lateral view. Fig. 35. Phyllocycla propinqua spec. nov. — Apical segments of abdomen, ® allotype, left lateral view. yellow. Pterostigma yellow, covering 6—7 cells. Two basal subcostal cross-veins in left front wing, one in other wings. Antenodal and postnodal cross-veins of first series 10 :17—18 : 10/12 :13—14:11 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth. Trigonal interspaces starting with a row of three cells against triangle followed by two rows of cells, seven cells long, in anterior row, in front wing, five cells long, in anterior row, in hind wing. Front wing with 4 rows of cells behind Cu2, hind wing with 5 rows of cells in the same area. Anal loop in hind wings two-celled. Hind wings with 5 paranal cells. Three postanal cells in right hind wing, four in left hind wing. Second anal interspace of hind wing starting with a row of two (paranal) cells against anal vein. J. BELLE: South American Gomphidae 235 Phyllocycla propinqua spec. nov. Fig. 35 Material. — Brazil: Santa Catarina, Nova Teutonia, 13.1.1949, 1 8; 20.X1.1950, 1 @ (holotype and allotype, respectively), (ML); Theresopolis, 1 , J. Michaelis; 1 & (paratypes, MNB); 1 & (paratype, coll. of author). In my 1970 paper on the South American Gomphidae this Brazilean species was considered no more than an eastern form of Ph. viridipleuris (Calvert). Since the female of Ph. viridipleuris is known from the type locality, Paraguay, I would con- sider the eastern form as a distinct species, closely related to Ph. viridipleuris, but readily recognizable from it by the following features: 1. Second lateral brown stripe of pterothorax partly developed and always represented by a distinct brown marking near subalar carina and at level of spiracle; third lateral brown stripe very weakly developed and on metepimeron generally represented by an ill-defined brownish stripe along femoral suture. In Ph. viridipleuris these stripes not developed at all. 2. Tibiae entirely blackish. In Ph. viridipleuris outer side of tibiae yellow. 3. Superior caudal appendages of male more slender and more acute at tip than in Ph. viridipleuris, and at about half length the inner surface nearer the upper edge there is only a low hump, marking the place of the small tubercle which is present in Ph. viridipleuris. 4. Lamellate expansion of ventro-tergal margins of eighth abdominal segment of female distinctly developed and much wider than that of the ninth abdominal segment. In Ph. viridipleuris very narrow and at most twice as wide as that of ninth abdominal segment. Male (holotype). -— Total length 52.5 mm; abdomen 40 mm (incl. caud. app.); hind wing 30.5 mm; costal edge of pterostigma in front wing 3.8 mm. Wings slightly brownish yellow at base. Venation dark brown but costa with an inconspicuous narrow yellow line on frontal margin. Pterostigma light brown, covering 41/,—51/, cells. Basal subcostal cross-vein present. Antenodal and postnodal cross-veins of first series 10 : 17—16 : 10/10 : 11—11 : 10 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth in left front wing, the fifth in other wings. All supratriangles, subtriangles and triangles two-celled, but subtriangle in hind wings open. Wings with 3 (front) and 4 (hind) rows of cells behind Cu2. Anal loop in left hind wing two-celled, in right hind wing one-celled. Five paranal cells and four postanal cells in each hind wing. Anal triangle in hind wing four-celled. Female (allotype). — Total length 49 mm; abdomen 37.5 mm; hind wing 30.5 mm; costal edge of pterostigma in front wing 3.6 mm. Venation of wings dark brown, including frontal margin of costa. Antenodal and postnodal cross-veins of first series 9 : 18—16 : 10/9 :14—13:11 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth in right wings, the seventh in left wings. All supratriangles, subtriangles and triangles two-celled, but sub- triangle in hind wings open. Wings with 3 (front) and 5 (hind) rows of cells behind Cu2. Anal loop in hind wings two-celled. Hind wings with 4 paranal cells and 4 postanal cells. Second anal interspace in hind wings starting with a single large (paranal) cell against anal vein, followed by two rows of cells. Remark: All other specimens from Brazil referred to Ph. viridipleuris in my 236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 1970 paper on the South American Gomphidae should also be considered paratypes of Ph. propinqua spec. nov. Phyllocycla pallida Belle, 1970 Material. — Uruguay: Lagas, source area, 1 g', Fruhstorfer (MNB). An old specimen, as clearly appears from the brownish wing membrane, differing from male holotype by the more obscure pale brown markings and more extensive darker markings. Face pale brownish but anteclypeus pale green. Antero-superior surface of the frons pale yellow. There is no distinct pale twin-spot on the prothorax. Inferior lateral angles of abdominal segment 10 somewhat produced inward. Anal loop two-celled in right hind wing and three-celled in the left. Anal triangle five-celled in both hind wings. Negomphoides undulatus (Needham, 1944) Material. — Brazil: State of Pará, Upper Rocana River, VI or VII.1918, 6 &', S. M. Klages (FSC). Negomphoides lieftincki Belle, 1970 Fig. 30, 31 Material. — Bolivia: Dept. Santa Cruz, Prov. Ichilo, pond stream locality, 18 km west of Buena Vista, 8 km west of San Carolos, 15.1II.1960, 1 © ; 17.111.1960, 1 dg; 20.111.1960, 1 9; Rio San Miguelito, 14 km east of Buena Vista, 22.111.1960, 1 8; Dept. La Paz, Prov. Nor Yungas, Canada, Naranjacada, a stream which crosses the road 4 km north of Caranavi, 11.1V.1960, 1 8; 12.1V.1960, 1 g'; Dept. La Paz, Prov. Nor Yungas, 1 km north of Caranavi, rapid mountain stream, 13.1V.1960, 2 3, all R. B. Cumming; El Cairo, 18.11.1960, 1 9; no locality, 12.V.1960, 1 g', 1 9; 15.V.1960, 6 &, 1 9; Dept. Cochabamba, Prov. Chapare (1600 m), El Palmar, no date, 1 9; Dept. Santa Cruz, Prov. Ichilo (400 m), Buena Vista, no date, 1 d'; 10.11.1960, 1 9, all R. Steinbach (FSC); Prov. Sara (600 m), no date, 1 9 ; no locality data, 1 g', both R. Steinbach (MNB). The markings on the dorsum of the pterothorax vary greatly in this species. All Bolivian representatives have a well-developed second pale antehumeral stripe im- mediately in front of the humeral suture. In the females the first pale antehumeral stripe is short and not connected with the pale area of the anterior mesothoracic “half collar’; in the males this stripe is longer and confluent with the pale area of the meso- thoracic “half collar”. Zonophora batesi Selys, 1869 Material. — Brazil: Para, Maranhao Aldeia Yavaruhu, Araçu, 50 km east of Caninde, 11—25.11.1966, 2 g', B. Malkin and José Celio Pinheiro. — Surinam: Maratakka (upper part), 5.111.1971, 1 8, D. C. Geijskes (ML). J. BELLE: South American Gomphidae 237. Zonophora calippus Selys, 1869 Fig. 39 Material. — Brazil: Para, Maranhao Aldeia Yavaruhu, Aracu, 50 km east of Caninde, 11-25.11.1966, 2 g', B. Malkin and José Celio Pinheiro. — Venezuela: District Anzoategui, between San Diego de Cabrutica and St. Cruz, 21.V.1936, 1 9. — Surinam: Kaboerikreek (upper part), 26.111.1971, 2 g'; 1.IV.1971, 1 g, D. C. Geijskes (ML). Zonophora spectabilis Campion, 1920 Fig. 36—38 Material. — Bolivia: Dept. Santa Cruz, Prov. Ichilo, Buena Vista (400 m), OM ¢ESG): Fig. 36—38. Zonophora spectabilis Campion, 9. 36, apical segments of abdomen, dorsal view, showing colour design; 37, diagram of pterothorax; 38, vulvar scale and ninth sternum, ventral view. Fig. 39. Zonophora calippus Selys, 2. Vulvar scale and ninth sternum, ventral view 238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 The female of this species differs greatly from Z. calippus, regarding the coloration of the abdominal segments 8 to 10, but the morphological differences of the vulvar scale are slight. Schmidt (1941) considered Z. spectabilis no more than a subspecies of Z. calippus. Z. spectabilis was described from Sapucay, Paraguay. Female. — Total length 50.5 mm; abdomen 37 mm (incl. caud. app. 2.8 mm); hind wing 38 mm; costal edge of pterostigma in front wing 4.7 mm, in hind wing 5 mm. Face predominantly yellow but free border of labrum blackish, depressed areas of postclypeus brownish, and vertical part of frons brownish above, becoming blackish in middle. Superior surface of frons yellow with a broad, blackish median band, which is connected with the blackish basal band. Vertex blackish, the depressed area behind and between the projecting postocellar ridges yellow. Occipital plate largely yellow, the lateral borders blackish. Rear of head brown but yellow behind occipital plate and on temporae. Labium and adjacent mouth parts yellow. Prothorax blackish, yellow on sides and on hind collar. Pterothorax blackish with yellow stripes, its pattern shaped as shown in Fig. 37. Dorsal side of femora yellow, becoming blackish to knees. Ventral side of anterior femur largely green. Tibiae, tarsi and claws blackish. Abdomen blackish with yellow markings. Markings of segments 7 to 10 shaped as shown in Fig. 36. Segment 3 to 6 with a broad yellow basal band interrupted on mid- dorsum. Segment 3 with a yellow lateral spot between basal band and posterior margin. Segment 1 and 2 yellow on middorsum, the sides largely yellow. Wings hyaline. Venation pale brownish but costa yellow from base to pterostigma. Pterostigma brownish yellow, surmounting 51/,—61/, cells. Basal subcostal cross-vein wanting in right hind wing, present in other wings. Each wing with two cubito-anal cross-veins in addition to inner side of subtriangle. Antenodal and postnodal cross-veins of first series 14 :19—20 : 14/12 :15—15 : 12 in front and hind wings, respectively. Second primary antenodal cross-vein the sixth in right hind wing, the seventh in other wings. Intermedian cross-veins 7—8/4—4 in front and hind wings, respectively. In each wing triangle two-celled, subtriangle open and supratriangle open. Hind wings with six paranal cells and five postanal cells, the sixth paranal cell is the first postanal cell. Trigonal interspace starting with a row of three cells against triangle, followed by two rows of cells, in front wings 1 (right) and 8 (left) cells long, in anterior row, in hind wings 3 cells long, in anterior row. Five rows of cells behind Cu2 in hind wings. Zonophora klugi Schmidt, 1941 Material. — Brazil: Upper Amazone, Sao Paulo de Olivenga, I.1923, 1 9, S. M. Klages (FSC). The present female has the supratriangle in the left hind wing two-celled, in the other wings open. The vulvar scale resembles greatly that of Z. calippus. Zonophora surinamensis Needham, 1944 Material. — Surinam: Tafelberg (foot), Geijskeskreek, 7.IX.1944, 1 g', L. Schmidt (ML). J. BELLE: South American Gomphidae 239 This is a new record for Surinam. The species was described from a single male taken in Brazil at Mapaoni, near the southern border of Surinam. The present male lacks abdominal segments 4 to 10. REFERENCES Belle, J., 1963. Dragon flies of the Genus Zonophora with special reference to its Surinam representatives. — Stud. Faun. Sur. 5: 60—69, pl. 3—4. ——, 1964. Surinam Dragon flies of the Genus Aphylla with a description of a new species. — Stud. Faun. Sur. 7: 22—35, fig. 5—14. ——,, 1966. Surinam Dragon-flies of the Agriogomphus complex of genera. — Stud. Faun. Sur. 8: 29—60, fig. 51—92, pl. 5—10. ——, 1966. Additional notes on some Dragon-flies of the genus Zonophora. — Stud. Faun. Sur. 8: 61—64, fig. 93—98, pl. 11. —, 1970. Studies on South American Gomphidae with special reference to the species from Surinam. — Stud. Faun. Sur. 11: 1—158, fig. 1—264, pl. 1—21. ——, 1970. On the Neotropical Gomphidae of Longinos Navás. — Tijdschr. v. Ent. 113 (6): 253—260, fig. 1—15. Calvert, P. P., 1901—1908. Odonata ir Biol. Centr. Amer., Neuropt.: xxx + 420 pp., pl. 2—8. (Gomphinae: June 1905). — 1903. On some American Gomphinae. — Ent. News 14: 183—191, pl. 8. ——, 1909. Contributions to a knowledge of the Odonata of the Neotropical Region, exclusive of Mexico and Central America. — Ann. Carnegie Mus. 6: 73—280, pl. 1—9. —, 1948. Odonata (Dragonflies) of Kartabo, Bartica District, British Guiana. — Zoologica, New York 33: 47—87, pl. 1—2. Campion, H., 1920. Some new or little known Gomphine Dragonflies from South America. — Ann. Mag. Nat. Hist. (9) 6: 130—141, pl. 6—7. Dias dos Santos, N., 1968. Notas sôbre o imago de Epigomphus paludosus Hagen in Selys, 1854. — Atas Soc. Biol. Rio de Janeiro 11 (4): 159—161, fig. 1—4. Fraser, F. C., 1947a. The Odonata of the Argentine Republic I. — Acte Zool. Lilloana 4: 427— 461, 4 fig. , 1947b. Notes on the Selysian types of Epigomphus paludosus Hagen in Selys. — Ann. Ent. Soc. Amer. 40: 672—676, 2 figs. Geijskes, D. C., 1964. The female sex of Cacus mungo, Gomphoides undulatus, Planiplax phoeni- cura, Planiplax arachne and Dythemis williamsoni. — Stud. Faun. Sur. 7: 36—47, fig. 15—21. Kirby, W. F., 1890. À synonymic Catalogue of Neuroptera Odonata or Dragonflies, London: IX + 202 pp. Navas, L., 1927. Insectos de la Argentina y Chile. — Rev. Estudios No. de Julio 1927: 22—25, fig. 9. , 1933. Insectos Suramericanos. — Rev. Acad. Madrid 29: 191—198, fig. 11—17. Needham, J. G., 1940. Studies on neotropical Gomphine dragonflies. — Trans. Amer. Ent. Soc. 65: 363—394, pl. 20—22. ——, 1943. Notes on some Gomphine dragonflies from Venezuela and Guatemala. — Bolet. Ent. Venezol. 2: 197—206, fig. on p. 200. —, 1944. Further studies on neotropical Gomphine dragonflies. — Trans. Amer. Ent. Soc. 69: 171—224, pl. 14—16. Ris, F., 1904. Odonaten in Hamb. Magalh. Sammelr. 7: 3—44, 14 fig. 1913. Neuer Beitrag zur Kenntnis der Odonatenfauna von Argentina. — Mém. Soc. Ent. Belg. 22: 55—102, 24 fig. Schmidt, E., 1941. Revision der Gattung Zonophora Selys. — D. Ent. Ztschr. 1941: 76—96, fig. 1—12. , 1952. Odonata nebst Bemerkungen über die Anomisma und Chalcopteryx des Amazonas- Gebiets. — Beitr. Faun. Perus 3: 207—256, fig. 1—13, tables 1—4. Selys Longchamps, E. de, 1854. Synopsis des Gomphines. — Bull. Acad. Belg. (2) 21: 23—112 (3—93 sep.). > 240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 5, 1972 —, 1859. Additions au synopsis des Gomphines. — Bull. Acad. Belg. (2) 7: 530—552 (3—26 sep.). ——, 1869. Secondes additions au synopsis des Gomphines. — Bull. Acad. Belg. (2) 28: 168—208 (5—45 sep.). ——, 1873. Troisièmes additions au synopsis des Gomphines. — Bull. Acad. Belg. (2) 35: 732—774 (5—46 sep.); Appendix (2) 36: 492—531 (47—87 sep.). —, 1878. Quatriémes additions au synopsis des Gomphines. — Bull. Acad. Belg. (2) 46: 408—698 (3—106 sep.). Selys Longchamps, E. de & H. A. Hagen, 1858. Monographie des Gomphines. — Mém. Soc. r. Sci. Liege 11: 257—720 (viii + 460 sep.), 23 pls, 5 tables. Williamson, E. B., 1916. A new Cyanogomphus. — Ent. News 28: 167—172, pl. 8—9. ——., 1918. Two interesting new Colombian Gomphines. — Occ. Pap. Mus. Zool. Univ. Mich. 52.114 pli? ———, 1919. Archaeogomphus, a new genus of dragon-flies. — Occ. Pap. Mus. Zool. Univ. Mich. 63: 1—8, pl. 1. ——-, 1920. A new Gomphine genus from British Guiana with a note on the classification of the subfamily. — Occ. Pap. Mus. Zool. Univ. Mich. 80: 1—12, pl. 1. No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff, 1968 — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.FI. 70.—. No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Ditsh & Uvarov and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. D.Fl. 25.—. No. 5. C. A. W. Jeekel, 1970. — Nomenclator generum et familiarum Diplopodorum, 412 pp. D.FI. 100.—. No. 6. J. G. 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TIJDSCHRIFT VOOR ENTOMOLOGIE Redactie (Editorial Board) . . . . . . A. Diakonoff, P. J. van Helsdingen, C. A. W. Jeekel, R. de Jong, J. Krikken, M. A. Lieftinck. Address... .°. . . . . . + Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden. The Journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. It appears in separate issues, forming an annual volume of 350—400 pages. Subscription rate: D.Fl. 62.50 per volume. MONOGRAFIEEN VAN DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING Redactie (Editorial Board) and address as for Tijdschrift voor Entomologie. The Monographs are intended for the publication of larger entomological papers on a single subject and will appear irregularly. Che following Monographs have been published: Hors série: F. T. Valck Lucassen et al., 1961. — Monographie du genre Lomaptera Gory & Percheron (Coleoptera, Cetoniidae), 299 pages, 739 figs., 2 pl., map. D.Fl. 60.—. No. 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. REMARKS ON MATSUMURAESES ISSIKI, WITH DESCRIPTIONS OF NEW SPECIES (LEPIDOPTERA, TORTRICIDAE, LASPEYRESIINI) by A. DIAKONOFF Rijksmuseum van Natuurlijke Historie, Leiden ABSTRACT Remarks on the value of specific characters in Matsumuraeses Issiki are made, four new species from Java and Nepal described and four other species recorded. Recent studies reveal more and more species that have to be attributed to the present natural genus; these are species either already described, but referred to wrong genera, or new species. Without doubt their number will continue to grow. The key species is ‘“Semasia” phaseoli Matsumura, 1900, described originally from Japan; for some time it has escaped attention of European students, so that the number of its synonyms is considerable. Still caution is required with synonymising already described species of Matsumuraeses. So Danilevsky & Kuznetsov (1968) pointed out that M. ochreocervina (Walsingham) is a distinct species; and I myself am claiming the same for M. metacritica (Meyrick). The important specific characters of the males are the peculiarities of the socii — even more so than the shape of the valva; but especially, the androconial modifications of the dorsum of the hind wings. The females, on the contrary, do not always show specific distinction of the genitalia and are often difficult to separate, the more so, because the colouring and markings of the two sexes may vary to a great extent. Matsumuraeses belongs to the same group as the large genus Cryptophlebia Walsing- ham, with a circumtropical distribution. This group is a specialized off-shoot of the tribe Laspeyresiini. The development of their structures apparently followed the same direction as that in lower Eucosmini or higher Olethreutini. I would prefer to place this group at the very end of the Laspeyresiini and not at the place of the second Palaearctic genus, as in the monograph of Danilevsky & Kuznetsov (1968). A part of the present material originates from a series of recent collecting expeditions by the Munich Museum (Zoologische Sammlung des Bayerischen Staates), Germany, to Nepal. This material is exceedingly rich and is in excellent condition. I am greatly indebted to that Museum, especially to Dr. W. Dierl, for entrusting me with the study of this material and for the permission to retain certain duplicates for the Leiden Museum collection. 241 242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 Matsumuraeses elpisma spec. nov. (Pl. 1 fig. 1—2, Pl. 4 fig. 1—2) 3 14 mm. Head pale fuscous; a small transverse crest on forehead. Palpus greyish- fuscous, with a pale base, median segment broadly triangular, terminal segment very short, exposed. Thorax pale fuscous. Abdomen pale fuscous-grey. Fore wing oblong, costa straight, a trifle concave in middle, apex prominent, sub- obtuse, termen sinuate above, rounded beneath, moderately oblique. Pale ochreous- fuscous, with a weak transverse striation (tips of scales darker), faintly suffused with greyish towards costa and basal third of dorsum. Faint purplish marbling over dorsal suffusion and sparse dotting along costa, with an indication of an oblique, dark trans- verse line before 1/6 (edge of basal patch); basal 4/5 of costa suffused with greyish, this suffusion dilated into a rather faint costal spot, oblong-oval, from 1/2 to 4/5; dark purple dots on closing vein: a pair in right, a single, in left wing; sparse irregular purple points, tending to form short, horizontal strigulae between veins on termen. Cilia whitish- fuscous, with indication of purple bars opposite terminal point. Hind wing rather light fuscous, glossy, with less than basal half pale grey; a deeper fuscous streak above and parallel to vein 1b. Cilia fuscous-whitish, dorsum normal. Male paratype darker, finely spotted, and costa throughout transversely strigulated with ferruginous, terminal fifth rather suffused with dark grey, discal dark points absent. Otherwise similar to holotype. ® 12.5 mm. Similar to the holotype, but costa not darker suffused, disc moderately suffused with fuscous-brownish from dorsum beyond base to costa, becoming darker posteriorly. Hind wing darker infuscated, with a darker apex and veins (except upper edge of cell, closing vein and basal half of vein 1b). Cilia whitish, with a slender fuscous sub- basal band, not reaching tornus. Male genitalia. Tegumen moderate, rather spherical, not narrowed along upper half, except towards apex. Socius, a minute, rounded and conical process, short-birstled over outer side. Tuba analis membraneous, moderately broad, gently narrowed, only extreme base darker. Valva long, sacculus oval, neck moderate, cucullus broadly semicircular. Aedeagus long, slender, curved, broadest at base, gradually narrowed. Cornuti, a sheaf of long, slender spines. Female genitalia. Sterigma simple, not modified. Ostium, a semiovoid cup, open above, moderately sclerotized. Lamella postvaginalis with two ear-like, small, minutely bristled patches. Ductus bursae narrow, colliculum, a small, weak cylindre. Cestum, a slender tube. Signa, moderate horns. Eighth sternite with a triangular sclerite in middle of posterior edge. East Java, Tengger Range, Tosari, 1770 m, bred from leaves of Crotalaria (Papilionaceae), 26.11.1940, 1 4, holotype, genit. slide 8731; 24.11.1940, 1 g', para- type; 14.11.1940, 1 9, allotype, genit. slide 8732 (A. Diakonoff). The same range, Nongkodjadar, 1300 m, at light, 1 9, 20.11.1940, genit. slide 8737; 1.1V.1940, 1 9, genit. slide 8736; 1 9, 6.IV.1940; 1 9, 31.1.1941 (A. M. R. Wegner). Apparently belongs to the Far Eastern M. monstruosana V. Kuznetsov group of species, but is distinct by the genitalia, especially the characteristic socii. A. DIAKONOFF: Remarks on Matsumuraeses 243 Matsumuraeses felix spec. nov. (PIN2 PIS) d 14.5 mm. Head light tawny-brownish with a faint purplish tinge. Palpus laterally with smoothly appressed scales; light tawny, median segment triangularly dilated, top darker brown, terminal segment concealed, pale tawny. Thorax tawny-brown, apical third and tips of tegulae strewn with purple, tip of thorax blackish-purple. Abdomen pale ochreous. Fore wing moderately broad, oblong, truncate, costa moderately curved throughout, apex obtusely pointed, almost rectangular, termen sinuate above, rounded below, vertical. Pale ochreous-tawny, markings dark purple. Basal patch to less than 1/4, edge formed by a series of closely set dots, angulate above middle, gently concave above and below angulation; this patch filled out by fine purplish dusting; central third of costa with three slender oblique lines not reaching cell, about equidistant, alternating with minute points; a conspicuous, dark purple spot under 2/3 of costa, formed by two parallel, short longitudinal streaks, interconnected by slight dusting, upper streak slightly continued on both ends above by slender lines; sparse marbling on second fifth of dorsum, connected with lower half of basal patch, strongly angulate posteriorly and extending along dorsum by sparse transverse dotting; a spot of faint dusting halfway between subcostal dark spot and termen; some three irregular vertical series of dots before and parallel to termen and a darker, spindle-shaped subterminal mark below apex, all these alternating with similar, very faint lines. Cilia pale tawny-fulvous, on dorsum becoming grey, basal fourth deep purple, becoming darker towards apex, paler on dorsum. Hind wing fuscous-bronze, posterior half dark grey in certain lights, basal third pale grey, becoming whitish at base. Cilia light fuscous-grey, with a darker subbasal band and a whitish basal line; cilia along dorsum slightly lengthened, otherwise normal. Male genitalia. Tegumen high, upper half narrowed. Socius, a small, obliquely rising oblong-oval plate, sparsely short-bristled, dilated tops slightly overlapping. Tuba analis wide, flat and narrowed above, with narrow edges sclerotized. Valva long, sacculus rhomboidal, lower angle at top projecting, pointed, lower edge moderately bristled; neck of valva narrow, cucullus broad, almost semicircular, top curved, projecting and rising. Aedeagus long, slender, strongly curved beyond base, gently dilated towards apical 1/3. (Cornuti missing.) East Java, northwestern slope of Mt. Ardjoeno, Tretes, 900 m, bred from leaves of Litsea (Lauraceae), 24.111.1940 (A. Diakonoff), 1 &, holotype, genit. slide 8730. Probably related to M. monstruosana V. Kuznetsov, from the Maritime District, Sibiria, and less closely, to M. melanaula (Meyrick), from India and Assam — now also recorded from Nepal. Matsumuraeses phaseoli (Matsumura) Semasia phaseoli Matsumura, July, 1900: 197 (& 2). — 1901: 23. — Yasuda, 1956: 16. Semasia elutana Kennel, August, 1900: 147, fig. 23. — 1916: 513, pl. 19 fig. 86. Eucosma trophiodes Meyrick, 1908: 613 (6 9 ). — Clarke, 1955: 319. Argyroploce trophiodes: Janse, 1917: 175. — Caradja, 1939: 12. Thiodia azukivora Matsumura, 1931: 1076, fig. — Obraztsov, 1952: 148. Lathronympha phaseoli: Issiki in Esaki, 1950: 480, fig. — Matsumura, 1953: 1, 2, 12, pl. 1 fig. 1; pl. 3 fig. 5, pl. 4 fig. 2. — Inoue, 1954: 95, no. 509. — Yasuda, 1956: 15, 17, figs. 1—4. Semasia azukivora: Yasuda, 1956: 15, 16. 244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 Figs 1—2. Male genitalia of Matsumuraeses species, holotypes. 1, M. xantholoba sp. n.; 2, M. tetra- morpha sp. n. A. DIAKONOFF : Remarks on Matsumuraeses 245 Matsumuraeses phaseoli: Issiki in Esaki, 1957: 57, pl. 8 figs. 257, 258. — Okano, 1959: 260, pl. 174 fig. 20. — Obraztsov, 1960: 134, figs. 86—91. — 1967: 32. — Danilevski & Kuznetsov, 1968: 237, figs. 127, 128, 131 (elutana Kenn., azukivora Mats., trophiodes Meyr., metacritica Meyr. synonyms). Matsumuraeses (Semasia) elutana: Kuznetsov, 1962: 346, fig. 7. Olethreutes trophiodes: Clarke, 1958: 559, pl. 278, figs. 1—1a (lectotype designated, wings, genitalia figured). The present synonymy might be neither correct nor complete, owing to the wrongly identified older material. Close future study and field observations, especially in Japan, will reveal the actual identity of this and allied Japanese species. The species is of moderate size, 17—18 mm, and with considerably varying colouring and markings, but usually pale ochreous, with a suffused dark fuscous streak along middle of costa and a darker grey suffusion along dorsum, dilated into an irregular patch at 1/3, sometimes also with a suffused discal stigma. The female usually is greyish- ochreous, with fine longitudinal pattern of thin lines more or less developed, sometimes with a diversely developed dark terminal shade. Hind wing in male pale ochreous, partly subhyaline, hyaline at base, with prismatic reflections, with dark fuscous, narrow marginal suffusion from costa to tornus, veins dark fuscous posteriorly; dorsum much thickened by closely appressed, modified and short scales, white, mixed with sparse jet-black scales and with long, pale ochreous- fuscous hairs. The male genitalia with moderately long, joined socii, with long, sideways-directed hairs, parted in middle; neck of valva less narrow than in other species, cucullus triangular and rather pointed, cornuti rather long and slender. West Java, Tjipanas, 1100 m, 1—10.X.1948 (A. Diakonoff), 1 4, genit. slide 8733. Buitenzorg, 250 m, bred from leaves of Tephrosia vogelii (Papilionaceae), 1 d'. — Mt. Gede-Pangrango, Tjibodas, 1400 m, IV.1949 (A. M. Neervoort), 1 G', genit. slide 8734, 2 & without abdomens. The same, IV—VI and XII.1949 (A. M. Neer- voort), 33 9, genit. slide 8735. — Lembang, 1100 m, 20.VIII.1922, 1 g', bred from leaves of “kedele” (Glycyne Max). East Java, Tengger Range, Nongkodjadar, 1400 m, 8, 11, 24.111.1940, 20.XII. 1940 (A. M. R. Wegner), 4 9, genit. slide 8738. Matsumuraeses xantholoba spec. nov. (Fig. 1) d 15-19 mm. Head purplish-fuscous. Palpus long, ascending close to face and exceeding vertex; median segment triangularly dilated by roughish scales above and beneath; terminal segment slender, smooth, pointed and exposed; greyish-fuscous, spotted in middle and on apex with purplish, terminal segment purple. Thorax rather dark grey- purple, tegulae light fulvous. Abdomen pale yellow, valva whitish, venter mixed with grey. Fore wing oblong-triangular, costa little curved, more so anteriorly, apex produced, obtusely pointed, termen strongly sinuate, little oblique. Whitish. Basal patch from 1/5 of costa to before middle of dorsum, edge with a round prominence below fold; this patch dark grey-olive, edged and dotted with purplish, lower half entirely deep purple, posterior edge and its prominence ferruginous; basal 3/4 of costa suffused with grey, transversely strigulated and dotted with purple strigulae, oblique and rather irregular; 246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 posterior half of costa with four oblong purple spots, first median, connected with the horizontal subcostal streak, three following costal spots becoming smaller posteriorly; a round purplish spot on middle of closing vein, an irregular group of paler, suffused spots, greyish and fulvous, below this, not quite reaching dorsum; a band of glossy grey and olive suffusion along edge of wing from below apex to end of dorsum, with minute purplish points between veins and three similar larger marks above tornus. Cilia deep purple, more or less suffused with black and ferruginous, with a white basal band. Hind wing grey-whitish, posterior half strongly infuscated from below costa, which is white, to vein 5 and in a suffused band along upper half of termen, veins 3-5 dark fuscous, cells between them thinner scaled, below these veins scaling glossy and more opaque, vein 1b dusted with black scales, sparsely strewn above this vein; dorsum forming an extended lobe, densely covered with thick, round, thin-stalked scales, from whitish below vein 1 b, turning rather bright yellow towards dorsal edge; under side of lobe with smooth and granulate, whitish-ochreous surface. Cilia fuscous with a white base and darker grey-fuscous subbasal band. Male genitalia. Resembling those of M. phaseoli, but with socii twice as long; valva with a narrower neck, triangularly dilated base of sacculus larger, as well as the group of bristly hairs in its centre; collicalum with more prominent and less obtuse angles. Nepal, Prov. No. 3 East, Jubing, 1600 m, 9.V.1964, 1 3, holotype, genit. slide 8569; the same locality, 6.V. and 11.V.1964, 2 g', genit. slide 8571 (W. Dierl), para- types. Superficially resembling M. phaseoli and also M. ochreocervina rather closely, but with the very characteristic yellowish dorsal lobe of the hind wing. Matsumuraeses metacritica (Meyrick) (Fig. 4) Eucosma metacritica Meyrick, 1922: 515 (4). — 1935, in Caradja & Meyrick: 35. — Clarke, 1955: 204. — Viette, 1957: 151. Allohermenias metacritica: Razowski, 1960: 385 (lectotype designated and genitalia 4 figured). Matsumuraeses metacritica: Danilevski & Kuznetsov, 1968: 237 (syn. of phaseoli Mats.). Distribution. S. China: Shanghai. Nepal, Prov. Nr. 2 East, Jiri, 2000 m, 2.IV.1964 (W. Dierl), 1 g', genit. slide 8570. It is a surprise to encounter this Southern Chinese species at an altitude of 2000 meters; however, there is no question about the identity of the single specimen available: although somewhat rubbed, the fore wing has retained most features, while the hind wing which is even more characteristic, is completely intact. The terse original description of the male hind wing may be elaborated as follows. Light grey, thinly scaled, with slight prismatic reflections; a moderate fuscous-grey suffusion along posterior third of upper half, more or less becoming darker in apex; veins distinctly but narrowly streaked with fuscous, except lower edge of cell from beyond base to angle and veins 2 and 3 throughout, which are dilated by blackish- fuscous dusting; dorsum corrugated, forming a fold above vein 1b, another below this vein, with a compressed fold between these, densely covered throughout on the upper side with closely appressed fuscous-grey hair-scales, cilia from end of vein 3 to tornus similarly infuscated, cilia along dorsum dense, elongate and whitish; a long pecten of fine pale grey hairs along first fold, a less extended but longer second pencil of similar hairs along base of second fold, extending on cell 1a and dorsum. Underside only with TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 PLAAT 1 Plate 1. Matsumuraeses elpisma sp. n., holotype. Fig. 1. Male genitalia. Fig. 2. Coremata; left posterior corema in place, right corema pushed outwards (caudad) A. DIAKONOFF : Remarks on Matsumuraeses TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 PLAAT 2 Plate 2. Matsumnraeses felix sp. n., male genitalia, holotype A. DIAKONOFF: Remarks on Matsumuraeses Sasaran unse UO SFAVULIYM] : JJONONVIC 'V eyewaroo ‘adAjopoy “u ‘ds xzjaf sasorumunsipw "€ Ed è € LVVTd TL6I ‘9 ‘TAV ‘SIT TIIA ‘HIDOIONOLNA YOOA LAIYHOSAIT S2S9PAHUNSIPN] UO SFAPUIM : JHONONVIC ‘V xınejndo> esang ‘7 ‘Sig erpepruag ‘1 “sry adMoyye è “U “ds rıusıdja Sasoranunsiry ‘% rye FP LVVId ELO ‘9 “TAV ‘CIT TATA ‘AIDOTONOLNY YOOA LATMHOSAIT], SAISAVAHUHSIPIAT UO SYAPUI ST : JAONONVIG 'V wnugis pue avsing sndioy ‘7 ‘Arq ‘ewS1193S ‘1 PIJ ‘etenu9$ ‘adore “u ‘ds pqdiourvuas sasavinunsyryy °C arid S LYVIg ZLOT ‘9 “Tav “CTT 1aaa “AIDOTOWOLNY HOOA LAINHOSAlLL Sasap4anWunsppy UO SFAPUIY] : AHIONOXVI( 'V 9 LVVId TL6I ‘9 ‘TAV ‘STI TATA ‘AIOOIONOLNY YOOA LAINHISA[TL A. DIAKONOFF: Remarks on Matsumuraeses 247 grey suffusion in cell 1c towards margin and thickened black posterior end of vein Ic, veins 2 and 3, and cell between these veins. Posterior tibia suffused with black. Male genitalia. Gnathos arms continuous, rather broad and distinct, coloured. Socti moderately long, a pending, oblong body with very long diverging bristles. Valva robust, sacculus triangularly prominent at top; cucullus subquadrate, with rounded angles, inner lower edge with a distinct angle, as far as this very densely short-bristled, along posterior edge also long-spined, spines hardly reaching middle of cucullus. A sketch and a brief description of the genitalia have been published by Razowski (l.c.), who attributed the species to Allohermenias and selected a lectotype. The sketch is very accurate, but the interpretation of the joined socii as a “haired plate of the gnathos”’ is incorrect. Matsumuraeses tetramorpha spec. nov. (Fig: 2, Pl. 5) d Q 18—22 mm. Head fuscous-brown. Palpus rather long, more than diameter of head projecting beyond face; median segment triangular, strongly dilated above and beneath with scales roughish along edges, terminal segment moderate, very slender, partly concealed, subobtuse; dark fuscous-grey. Thorax purplish, strewn with sparse white scales, tegula tawny-fulvous, collar fuscous-brown. Abdomen glossy pale ochreous, tinged fuscous except apex, venter tinged grey. Fore wing oblong-triangular, moderately dilated and broadest at 3/4; costa curved throughout, more so at base, apex pointed and slightly prominent, termen considerably sinuate and oblique. White, along costa and towards tornus finely dark-striated (dark bases of scales). Costa, as far as upper edge of cell, marbled with fulvous, with a fulvous patch at base and six transverse marks, anterior two wedge-shaped, continued to cell by slender, oblique lines, third and fourth marks similar, but much broader, fifth and sixth subquadrate, without appendages; all these costal marks alternating with slender lines; extreme edge of costa between markings finely striated; from about third to fourth fifth of disc, from below costa to as far as above middle of dorsum, with a pattern of some five more or less suffused series of partly interconnected irregular spots, rows tolerably parallel to termen, spots interconnected above, becoming more separate and larger down- wards and turning from bright tawny-fulvous to pale olive; dorsum suffused with fuscous-brown, from beyond base cut by seven white spots, becoming wider posteriorly; fine fuscous-brown dusting towards base, over centre of disc; a well-defined, straight and slightly inwards-oblique series of six black points above tornus, upper dot below vein 8; these dots slightly mixed with tawny-olive and followed by greyish suffusion; two minute points just before termen below apex, indicating a second series of black points; a jet-black raised mark along lower part of closing vein, followed by dark dusting. Cilia dark purple, with a strong gloss, around tornus more or less mixed with white. Hind wing glossy pale fuscous, dorsum normal, tornus not in any way thickened, only slightly concave on the underside, hairs almost normal, except longer pencils of loose fine hairs along the bases of all the three anal veins on upper side and less densely spread over the basal half of cell 1a. Cilia glossy fuscous-whitish, with a light grey narrow subbasal band. 248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 Except this pattern of colouring and markings there are three other, more or less interconnected by intermediates, viz.: (1) Reduced markings of the above described pattern, with ground colour pale grey, costal markings reduced to a diverse number of fuscous oblique streaks, dorsum with a fuscous suffusion, discal black spot distinct, but the preterminal row of black points diversely reduced. Both males and females. Khunjung, Bujan. (2) Melanic pattern: deep purplish-fuscous, with all veins streaked with whitish and posterior half of costa with five diversely developed whitish spots. Both males and females. Khunjung. (3) Indifferent pattern: whole wing densely suffused with diverse shades of grey, purplish-grey to pale grey, sometimes with two rows of dark preterminal dots, some- times with a diversely developed pale spot between cell and termen. Most females belong to this colour form. Khumdzung. In females often there is a shadow along dorsum and four faint darker dots along costa posteriorly. Male genitalia very similar to those of M. phaseoli Matsumura and differing only by the following minor particulars: socii rather smaller and considerably less vigorously bristled, with shorter bristles; valva with a narrower neck, cucullus more triangular, its lower point more produced; sacculus more rounded. Cornuti distinctly longer. Female genitalia are similar to those of M. phaseoli. Nepal, Khumbu, Tangpoche, 4000 m, 10.VII.1962 (G. Ebert & H. Falkner), 1 9, paratype. — Pangpoche, 4000 m, 4 and 10.VII.1964 (W. Dierl), 2 9, paratypes. — Khumbu, Khumdzung, 3900 m, 15—25.VII.1962 (G. Ebert & H. Falkner), 5 8,2 9, genit. slide 8568, paratypes. — Prov. No. 3 East, Kumjung 3800 m, 25, 27 and 30.VI, 1, 12, 14 and 18.VII.1964 (W. Dierl), 75 &, genit. slide 8564, holotype, 8565, para- types; 11 ©, allotype with genit. slides 8573; 8566, 8568, paratypes. — Prov. No. 3 East, Bujan, Dudh Kosi Tal, 2900 m, 21.VI.1964 and 18—19.VII.1964 (W. Dierl), 8 d, 1 2, paratypes. — Prov. No. 3 East, Junbesi, 2750 m, 25—31.VII.1964 (W. Dierl), 1 g. — Prov. No. 2 East, Jiri, 2000 m, 13.VIII.1964 (W. Dierl), 1 9, genit. slide 8567. All paratypes. — Kathmandu, 1400 m, 23.VII.1962 (G. Ebert & H. Falk- ner), 1 9, paratype. The extremes of diverse types of colouring and markings of this species are so striking that doubts may arise as to their conspecificity. However, they are connected by inter- mediates and have the same robust build and the shape of the pointed fore wings, while a scrutiny of the male genitalia did not reveal any differences. This species of high mountains apparently is closely allied with M. phaseoli Matsumura, but differs at once by a simple dorsum and tornus in the male, not thickened with modified dense scales, as in that species. Matsumuraeses melanaula (Meyrick) (Fig. 3) Eucosma melanaula Meyrick, 1916: 17 (& 2). — Fletcher, 1920: 49, pls. 1—2. — Clarke, 1955: 199. — 1958: 372, pl. 185 figs. 1—1a (lectotype designated and figured). Matsumuraeses melanaula: Issiki in Esaki c.s., 1957: 57 (comb. nov.). — Obraztsov, 1967: 34. — Danilevski & Kuznetsov, 1968: 67. Eucosma melaneura Yasuda, 1956 (lapsus): 16, 17. Distribution. Assam, Bengal, South India. Food plants. Cajanus, Phaseolus (Papilionaceae). The species may be redescribed as follows. A. DIAKONOFF : Remarks on Matsumuraeses 249 (Ce INS a IN is Fig. 3—4. Male genitalia of Matsumuraeses. 3, M. melanaula (Meyr.), slide 8562; 4, M. metacri- tica (Meyr.), slide 8570 3 250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 d 14—16 mm. Head and palpus rather dark fuscous-grey, palpus paler ochreous on basal 2/3 of median segment. Thorax light ochreous-grey, shoulder and dorsum, except anteriorly rather suffused dark grey. Posterior tibia smoothly scaled, pale ochreous. Abdomen whitish, tinged pale fuscous-grey. Fore wing oblong-subtruncate, moderately dilated, costa curved anteriorly, almost straight posteriorly, apex obtusely pointed, scarcely prominent, termen moderately sinuate above, rounded below, little oblique. Pale ochreous faintly dusted with dark fuscous and with a few black specks here and there (paratype rather evenly suffused with fuscous- grey); costa with a series of numerous short, purplish comma-like marks, alternating with minute strigulae and gradually becoming longer posteriorly; the larger marks continued by oblique tawny suffused faint streaks to upper edge of cell, each streak dilated there into an irregular small spot, so as to form a series of spots below and parallel to costa, from beyond base to about 3/4; a fragment of a larger tawny-fulvous costal spot at 1/3; together with an inwards-oblique bar on 1/3 of dorsum indicating edge of basal patch; the dorsal bar reaching fold, with top rounded; a large semicircular- erected tawny-fulvous patch on dorsum beyond middle, gently inwards-oblique, top to lower angle of cell; a suffused small black horizontal striga beyond cell in middle of disc; dorsum from beyond base irregularly clouded with purplish, not reaching beyond halfway fold, except in tornus where clouding becomes black, less regular and almost reaches vein 4; ocelloid spot ill-defined, edges hardly perceptible, grey, centre with three black marks; some tawny striae along termen. Cilia pale ochreous, densely marbled with purplish, with minute snow-white tips, a broad basal fulvous streak. Hind wing fuscous-grey, rather thinly scaled, narrow bases of scales being whitish- hyaline, darker suffused along veins, paler towards costa anteriorly, dorsum with a golden gloss, a streak of dull jet-black scales above vein 1b on underside, showing through as a grey streak on upper side. Cilia light golden-ochreous with a fuscous subbasal line, cilia along dorsum elongate and slightly curly, otherwise dorsum normal. Male genitalia. Top of tegumen with a large semicircular, laterally compressed naked prominence, socii absent. Gnathos represented by two sinuate, submembraneous lobes. Valva moderately constricted, sacculus subrhomboidal, with an oblique postmedian series of hairs, lower angle distinct but obtuse; cucullus semioval, lower edge and top strongly rounded, top rather narrow. Aedeagus very slender, sinuate, cornuti, three deciduous spines with subterminal bases and scattered minute spines. Nepal, Kathmandu Valley, Godavari, 1600—1800 m, 18.1X.1967 (Dierl-Forster- Schacht), 1 &, genit. slide 8501. — Kathmandu, Chauni, 1400 m, 5.IX.1967 (Dierl- Schacht), 1 3, genit. slide 8562, figured in this paper. The paratype is suffused with grey, strongly obscuring other markings. Characteristic are the narrow and the broad inwards-oblique dorsal spots, and the black androconial patch in the anal region of the underside of hind wing. Matsumuraeses ochreocervina (Walsingham) (Pl. 6 fig. 1—2) Eucelis ochreocervina Walsingham, 1900: 407 (& 2). — Yasuda, 1956: 16. — Obraztsov, 1960: 134 (syn. of phaseoli Mats.). — 1967: 33 (syn. of phaseoli). Laspeyresia ochreocervina: Inoue, 1954: 482. Matsumuraeses ochreocervina: Danilevski & Kuznetsov, 1968: 239, figs. 129, 132 (distinct species, genit. 2 figured, biology, distribution). A. DIAKONOFF: Remarks on Matsumuraeses 251 Distribution. Japan, Mongolia, Sibiria, Far East. Food plant. Astragalus membranaceus (Papilionaceae). The species is larger and more robust than M. phaseoli: 23—24 mm, ground colour whitish-ochreous, a dark grey suffused streak along costa not reaching apex nor base, a light brown line along dorsum, dilated at 1/3 into a rounded spot reaching 1/3 across wing and a fuscous-brownish suffusion over termen as far as cell, well defined edge along closing vein. In female no. 8574 this pattern is somewhat obscured by dark dotting over basal 1/3 of wing and terminal suffusion lacking. Hind wing light grey-fuscous, darker and more opaque on apical third, paler on dorsum; dorsum normal, without modified androconial scales, but cilia much lengthened, basal half of cell 1a densely long-haired (with longer and denser hairs than in M. tetra- morpha sp. n.). Hind wing in female similar. Male genitalia are well figured by Danilevski & Kuznetsov, 1968, fig. 129; they are characteristic by the socii shaped as a small rounded tumescence with moderately dense and not very long thin bristles; these socii are prominent upon a transversely flattened top of tegumen; the neck of valva is narrow, the angles of cucullus moderately rounded (less so than in fig. 129, cited above), cornuti slender and long. Female genitalia, cf. Danilevski & Kuznetsov, fig. 132, with lamella postvaginalis large, sclerotized, moderately narrowed downwards and deeply notched at base, actually little differing from those in phaseoli and tetramorpha. Colliculum moderate, without structures, rather wide, cestum distinct (not indicated in fig. 132). Signa large, almost of equal size and shape. Nepal, Prov. No. 3 East, Khumjung, 3800 m, 18.VI.1964, 1 3, genit. slide 8572; the same, 14.VII.1964, 1 g', 1 9, genit. slide 8574 (W. Dierl). Postscript After having sent this paper to the printers’, I received from Professor Dr. T. Yasuda information on Matsumura’s Japanese species of the genus Matsumuraeses and on their types, for which I wish to express my gratitude in this place. Prof. Dr. Yasuda, it appears, is preparing a paper on the biology of these Japanese species, which, I hope, will, once and for all, put an end to the misconceptions about their true identity. REFERENCES Caradja, A., 1939. — Materialien zu einer Microlepidopterenfauna des Mienshan, Provinz Shansi, China. — Iris: 1—32. Clarke, J. F. Gates, 1955. — Catalogue of the type specimens of Microlepidoptera in the British Museum (Natural History) described by Edward Meyrick, 1: I—VII, 1—312, fig. 1, frontispiece, pls, 1—4. — 1958, loc. cit. 3: 1—600, pls. 1—298. London. Danilevski, A. S. & V. I. Kuznetsov, 1968. — Fauna of the U.S.S.R., new ser., no. 98. Lepidoptera 5 (1), Leafrollers, Tortricidae, tribe Laspeyresiini; 1-—635, figs. 1—469 (in Russian). Leningrad. Fletcher, T. B., 1921. — Life-histories of Indian Insects, Microlepidoptera. — Mem. Dept. Agric. India, Ent. 11: I—IX, 1—246. Pusa. Inoue, H., 1954. — Check list of the Lepidoptera of Japan. 1. Micropterygidae — Phaloniidae: I—XIII, 1—44. Tokyo. Issiki, S. T., in T. Esaki c.s, 1950. , 1957. — Icones heterocerorum japonicorum in coloribus naturalibus, 21: 52—89. Osaka. 252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 6, 1972 Kennel, J., VIII.1900. — Neue palaearktische Tortriciden, nebst Bemerkungen über einige bereits beschriebene Arten. — Iris 13: 124—159. , 1916. — Die palaearktischen Tortriciden. — Zoologica 21: 397—546 + 8 pp., pls. 17—20. Kuroko, 1957. — Enumeratio insectorum montis Hikosan, Lep. 1: 10 pp. Kuznetsov, V. I., 1962. — New species of Leaf-rollers (Lepidoptera, Tortricidae) from the Far East. — Trudy Zool. Inst. Acad. Sci. U.S.S.R. 30: 337—352 (in Russian). Matsumura, S., VII.1900. — Neue japanische Microlepidopteren. — Ent. Nachrichten 26: 193— 199. , 1931. — 6000 illustrated Insects of Japan-Empire: 1—1497. Tokyo. Meyrick, E., 1908. — Descriptions of Indian Micro-Lepidoptera. — J. Bombay Nat. Hist. Soc. 18: 613— 638. , 1916. — Exotic Microlepidoptera 2 (1): 1—32. , 1922. — Loc. cit. 2 (17): 513—544. Meyrick, E. in A. Caradja & E. Meyrick, 1935. — Materialien zu einer Microlepidopteren-Fauna der chinesischen Provinzen Kiangsu, Chekiang und Hunan: 1—96. Berlin. Obraztsov, N. S., 1952. — Thiodia Hb. as not a North American genus. — Ent. News, 63: 145— 149, , 1960. — Die Gattungen der palaearktischen Tortricidae. II. Die Unterfamilie Olethreu- tinae. 3. Teil. — Tijdschr. Ent. 103: 111—143, figs. 64—106, pls. 11—13. —, 1967. — Loc. cit. III. Addenda und Corrigenda 2. — Tijdschr. Ent. 110: 13—36, pls. 1—2, Okano, M., 1959. — Phaloniidae, Olethreutidae, Tortricidae, in Iconographia insectorum japonico- rum colore naturali edita. 1 (Lepidoptera): 259—268. Tokyo. Razowski, J., 1960. — The type specimens of the species of some Tortricidae (Lepidoptera). — Acta Zool. Cracoviensia 16: 463—543, figs. 1—176. Viette, P,, 1952. — Les types des Tordeuses de Meyrick appartenant au Muséum de Paris. — Bull. Soc. ent. France, 57: 148—152. Walsingham, Lord, 1900. — Asiatic Tortricidae. — Ann. Mag. Nat. Hist. ser. 7, 5: 368—386, 451—469, 481—490; 6: 121—137, 234243, 333341, 401—409, 429448. Yasuda, T., 1956. — Revision of two moths of Lathronympha phaseoli Mats. and Semasia azukivora Mats. — Bull. Ent. Lab. Coll. Agric., Bur. Ent. Plant Quar. Osaka Pref. 2: 15—18. No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp. map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff, 1968 — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae)). 350 pp. en 98 pl. D.FI. 70.—. No. 4. F. 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Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. FURTHER STUDIES ON OLD WORLD MELECTINE BEES, WITH STRAY NOTES ON THEIR DISTRIBUTION AND HOST RELATIONSHIPS (HYMENOPTERA, ANTHOPHORIDAE) by M. A. LIEFTINCK Rhenen, Netherlands ABSTRACT with 2 plates, 57 text-figures, 1 table and 2 maps The present paper is a revisionary study of the smaller genera of parasitic anthophorid bees of the tribe Melectini occurring in the Eastern Hemisphere. The eight Old World genera pres- ently recognized are redefined and arranged in a key, which includes the diagnoses of two new genera of Ethiopian bees, Afromelecta and Acanthomelecta, each with a single previously described and named species. With the exception of the large and polymorphous genus Melecta Latr., which will be dealt with in a separate article, three new species and 14 earlier described taxa are analysed and, where necessary, redescribed and illustrated. Keys for the identification of species are given for all genera. New species described are: Protomelissa tricolor, Pseudomelecta atroalba and chalybeia. New synonymies established are: Callomelecta Ckll. (= Protomelissa Friese), Antho- phora sauteri Friese (= Protomelissa formosana (Ckll.)), Melecta glasunowi Mor. (= Paracrocisa kuschakewiczi (Radoszk.)), and Melecta spinosa Friese (= Acanthomelecta bicuspis (Stadelmann) ). The taxonomic treatment is preceded by general notes on the biology of Melectini throughout the world, followed by a preliminary list of concurrent Melectini and Anthophorini, whose parasite-host relationships have been either authenticated or suspected. CONTENTS WOtrOCUCHODMPEME, Speen ike isles er he Rf aoe e fom Ais on et Se oa 1254 Acknowledgements . . Sls Sp Wife (Ss, tea et we, eh Be ne ys 254 General biology of the Melectini 20 nh, GEE ee AE Host relationships . . . Ro IE a A an ten 290 Distribution of regional genera wee species eect Neh ere al ee Be ee men es 1260) Key to the Melectine genera of the Eastern Hemisphere. . . . . . . . . . . . . . 260 Discussion of genera and species (exclusive of Melecta Latr, 1802) . . . . . . . . . 266 ipyreusmpanzer, 1806... =. a Laos à nn à à à Fa à à a à à 266 Eynavlouskia (Popov, 1953 … €. ot et tene a à à à à de à s « + + 267 Protomelissan Briese, JOTA UT M UM ea is à GS à à à ee + ss à 268 Key to the species . . À ID, Pseudomelecta Radoszkowski, 1865 . DRE e Lee: Gey de asa cu Co) Gt GR 1202 Ikey.to the Species. = … … … e ee on En ee ee e 285 Paracrocisa Altken, 1937 … … è . 000 u 6 ie à à e es : … u 299 Keyrtoe the Species 4) to 2 00 eet Re el en en 301 Afromelecta gen. nov. . … 2: . . . . . . . . . . . 309 Acanibomelecias2ens NOV. «2 seis we Pb «ee ien dn ta a sn 316 References . . a ERR a 320 Alphabetical index È. NAMES koa A OEE 253 254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 INTRODUCTION The present account is one of the last in a series of revisional papers dealing with Old World Melectini, started by the writer in 1944 (Lieftinck, 1944—1969), and may be regarded as a continuation thereof. It aims at a critical study of all smaller and least known genera assigned to the tribe, some members of which were tentatively analysed already by Popov (1955). The components of this parasitic group are presently under- stood to be most closely related to their own hosts, i.e. the pollen-collecting bees of the tribe Anthophorini, with which they form one subfamily, Anthophorinae. A critical review of the dominant Holarctic genus Melecta Latr., with its numerous species in the Mediterranean basin and Eurasia, has not yet been given, but is in course of preparation and will, it is hoped, be published in a forthcoming final part of the series. ACKNOWLEDGEMENTS In the following list I have employed a series of symbols indicating the present location of the material studied, followed by the names of all colleagues, who have generously allowed me to study the specimens under their care. I wish, therefore, to express my sincere thanks to all who have helped me to make this survey as complete as possible. An important phase of this research has been the study of type specimens in various museum and private collections. I am especially indebted to Dr. H. Bytinski-Salz, for- merly of Tel-Aviv University, for his interest and friendly support, now several years ago, in the location of melectine types in the Museum fiir Naturkunde at Berlin. Dr. E. Königsmann, of the same institution, and Professor Dr. M. N. Nikolskaja, of the Leningrad Museum, have provided considerable facilities for the examination of critical specimens on the occasion of my scanning the Leningrad and Berlin collections of Hymenoptera, during August 1968 and the autumn of 1970, respectively. Freedom of access to all component parts of these and many other important collections has frequently resulted in the discovery of individual specimens that were temporarily lost, but could now be restudied. AMNH — American Museum of Natural History, New York (J. G. Rozen, Jr.) BM — British Museum (Natural History), London (I. H. H. Yarrow) BRNO — Moravské Museum (Entomological Department of the Moravian Museum), Brno, CSSR (Jaroslav Stehlik) CBS — H. Bytinski-Salz collection, Tel-Aviv CK — M. Kocourek collection, Vyskov, CSSR CW — K. Warncke collection, Dachau IEM — Instituto Espanol de Entomologia, Madrid (+ G. Ceballos and Senia Elvira Mingo) IZK — Instytut Zoologiczny (Institute of Systematic and Experimental Zoology), Polish Acad. Sci., Krakow (Miss M. Dylewska) MBUD — Magyar Nemzeti Muzeum (Hungarian National Museum), Budapest (J. ed) MHW — Martin Luther Universität, Halle-Wittenberg (R. Piechocki) ML — Rijksmuseum van Natuurlijke Historie, Leiden MNB — Museum für Naturkunde an der Humboldt Universität, Berlin (E. Königs- mann) M. A. LIEFTINCK: Old World Melectine bees 255 MP — Muséum National d'Histoire Naturelle, Paris (Melle S. Kelner-Pillault) MZUC — Museu e Laboratorio Zoologico, Universidade Coimbra (M. de A. Diniz) NMW — Naturhistorisches Museum, Wien (Max Fischer) NRS — Naturhistoriska Riksmuseum, Stockholm (S. Erlandsson) TMJ — Transvaal Museum, Pretoria (H. N. Empey and C. K. Brain) USNM — National Museum of Natural History, Washington, D.C. ZIL — Zoological Institute Acad. Sci. USSR, Leningrad (} A. Ponomareva, M. N. Nikolskaja, and V. I. Tobias) ZMM — Zoological Museum, Moskow (A. N. Zhelokhovtsev) GENERAL BIOLOGY OF THE MELECTINI I have taken the present opportunity to summarize briefly the most remarkable facts pertaining to the life history of the Melectini on the whole, as contained in the literature on the subject. Early stages Among the authors who have supplied first-hand knowledge of this kind the following deserve special attention. The most instructive and well-illustrated account dealing with the larval stages of the melectine Anthophoridae generally, is the one offered by J. G. Rozen (1969a). This important contribution to our knowledge of the tribal phylogeny was followed by an interesting memoir published by the same author on the biology of a new Thyreus, parasitic on a species of Anthophora, described from the Cape Province in South Africa (Rozen, 1969b). The larvae and pupae of the Australian Thyreus caeruleopunctatus (Blanch.) and /»gzbris (Smith), inquilines in the nests of Amegilla pulchra (Smith), were described by Cardale (1968b). Linsley & MacSwain (1942) reported some of the features of the biology of the Nearctic Xero- melecta (Melectomorpha) californica (Cresson) as a parasite of Anthophora linsleyi Tim- berlake (Hurd & Linsley, 1951). A complete description and figures of the larva of the last-mentioned cuckoo-bee are to be found in a paper by Michener (1953). Giordani-Soika (1936) briefly discussed the first instar larva of Melecta (M.) luctuosa (Scop.) in Italy. Torchio & Youssef (1968) published observations on the biology of the Nearctic Anthophora (Micranthophora) flexipes Cresson, and the immature stages of its parasite Zacosmia maculata (Cresson), with notes on the first instar larva of Melecta (M.) pacifica Cresson. Descriptions of the pupa of Melecta are to be found in papers by Semichon (1922) and Porter (1951). Host relationships The object of the Table here presented is to bring together our present knowledge of the inquilinic relation of various Melectini with their hosts throughout the world. This global survey does not pretend to be in any way complete, but gives an impression of the lacunae and may stimulate further research in this field. I have included in this Table also some of my own field observations made in the Oriental tropics, on various Amegilla presumably serving as the proper hosts of the species of Thyreus listed. However, I have been unable to definitely establish these specific relationships, owing to the absence of the nesting sites of the host bees involved. Hence in these and many TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 256 DLL Le VSN 392 A PUE[JOZIAS ULI] !Aurunan) odomx ANA EN) adomy Key] vIsy ‘M fodomg 'S “y's ‘aouraorg ode) (Lorsurr 2044218 “WY qns ‘VEFGI) Loysurr (6961) uazoy ‘ (1661) Aaysury 29 pimH (0161) Jaussaxy-Aar,y (SZ61 ‘9061) vooras ‘(T681) 2211 (ZZ61 ‘voyorwos ‘ZI6T ‘voxgry 'F°) ‘Arp “one (Z961) TPM (ZIGI ‘ua ‘39) ‘Arp one (9£61) Eros ‘9 (6961 ‘9961) PunJarT ‘(6S61) aodog (9761) sunvig ‘dsqns ‘yarwg vasaurs sesdosoqgduy UOSS2J7) ISprempa eroydoyuy CIID vided py 9I[MIgI SISTEHAMI *V (CT) wnyoaza9e eroydowuy LOSS9IJ vursan piogdogsuy CT) esnjar “y pur (‘zueg) sieansar eroydoyuy (nus) v/2,0u0z *H pur (vjourds) visa vpodosqvy smog vazsaf (i) vpodosgrH umouyun UOSSAID 991904091 (W) ‘W ‘TA ernjpea eyeredas (WW) ‘IN CD) wepund (Ww) ‘Ww Sajsur] vonurpe vorftvd (Ww) ‘Ww doos) vsomony (Ww) “IN [PIET V129/9W ‘dsqns"2 (osaray) 2449590 ‘A (quus) vuviounl “7 Aodod vrysaojavdugq (uozowe) qoaqnf ‘y “AOU ‘U9$ #/22J21404Îp7 (uuewjopes) sdsu1q ‘y ‘AOU ‘U9$ PIIIJIULOGIUPI LY ergeH Joymy Guroydoguy) 10H (1U199]PN) onsvsed ‘addy prog ur pauud are suonerosse paysi[quisg "tursoydoyyuy pue un SNOTITA UdIM}Oq SUONE]9I ysoy-azisvivd Surmoys ojgeL 257 M. A. LIEFTINCK : Old World Melectine bees eısy ‘1JU97) eısy U) VIEWING 'S NN BIDON weis UEAMIEI, LISV "UA VAE L Le) (SS61) aodog (6S6I1) Aodog (9961 ‘FF6T) PUnPrT (9961 pr61) Punzarı Cdo ‘nq) spunyor] (6761) 11712409 Cdo ‘nq) PURA] . (‘10W zaounspjd “dq qus ‘C¢61) Aodog (6€61) Aafsur] ‘yay z6ysjos vsogdogiuy “JON zuasuvg (T's) vaogdogzuy umouyun umouyun UMOUXUN (37217) suorvqdur vpodoigdva umouyun umouyun umouyun CTI) vavo vpodoagdvig ‘ds vpodorgdeig zo “ds (¢°Sqns) vpodoigvH ‘TD szsuodorns (¢°Sqns) vpodosgrH ‘vads (c‘#qns) vpodorgr umouyun “JON zodwas (vijisauvirg) vaogdoqiuy umouyun ‘ds s:sdosogdug i (som) szuuodissuf ‘q Cwszoag) vgzursvıp ‘q ‘aou ‘dads vzagkjvgs ‘d (ION) sadzy ‘a ‘aou ‘dads vq/vosgo vd “yzSOpry Mjsajaulopnasg (37017) vinsodpna ‘q ‘nou “dads 40709244 ‘d CIPID) Mangarpuad “qd 2SIT I SUI9SAPU ‘d CID) psorpisun “d (weySurg) vurlvppurg ‘a (TP) 20podosgrg ‘à (TO) pursowmsof “qd SOIT PSS7]2U0104d USS VAI: A I (‘yzsopey) sz27mayvqosny ‘gd (Pwsng) 12920113 ‘4 UONJIV 2220490404 UOSS2IJ uspsompa (snuw0199]W) IV TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 258 LOGE puejsuoon®) ‘SJ UOWOJOS svoonjoW °N ASN :efensuy 'S Asupis ‘pursusanê) aauraoid ade vaef fxeSunH adomg EBT eÁejewopur ‘] ouedug vÁejewopur BEHOBEN puesu2anê) (¢) erueumy vI}LVUINS vaef vende H (46561) spunzorT (28961) afepıe} paysıpgndun paysqqndun (SEGT) quawdry (2961) PURJIT ‘(q-e8961) STepıe) (q6961) u2z0y (961) \PUNJAT (8961) IPUNJAIT (661) 2H9YPIIS ‘(L761) JJopsig ‘(0Z61) vonag (Z961) NPURJAT (T961) PURPIT (Z961 PunJor] (‘psrog) vwososdwr] ‘49 qns ‘SE6T) yuswfey (q-r8961) arepseD (8661) winı (Z961) PunprI (z961) PUNPTT Joyny ‘dsqns (yyrwg) surdara vyedaup (yng) 2499724 eBay (TRIO) suardvs vou CIID) sesuaiwusay va ‘vads ‘y (ynws szuuofıgwog i) ‘ds epodoresy (x 38211 x eueisuneig vioydoquy (39917) 290901 y ‘(‘3J91T) veuwosof vyisaup (doy) ruadiqn y CITA) wepsezapenb yypsowy same 29 (III) zsmaspur vypsoup ‘dsqns (I) zsmaapur va sane 9 (IND) Zsmaupur v]pdauy (guawsey) ssuolounu py Crus) vaysind epSoury Crop) orages oyprdauy (35917) svqdopo vx ou CAT) 190928 ‘py ‘(IHAT) ovupuviaks “yy AJOUT 270.09 v]pd AU Guuoydoquy) 50H CID) soma — CD sumpum — CIPD) meus — Jar] suispdso sujnpyiu Crus) staqn3ny uazoy PPUNJI] Very 4941 CITI) snorwornsry ‘dsqns ‘2 (:pey) szruadoppung ‘Yor Stasau ‘I ‘dsqns ‘2 (asarı]) swazuojlas Cyouejg) snuoundospnrsrs Copy) surfe (Sax) 577044504 vp (asarag) szipunuopgp 'v Jazued smaulgL ‘mmm (UTP) anseIeg ‘mn 259 M. A. LiEFTINCK : Old World Melectine bees wsn qn "VST Sea Yon nu) vIUIOJITE) vruIoge) e1UJO JIE) ‘WS ‘HU ‘wIuUIoJe) adomg vof N vIPUI ‘S adomy epnsuruag Are VIJEAEH vaef ‘] uoquy (26961) uszoy £(8961) Jassnox 29 OIYDIOL, (1561) SPIH ‘Das ‘KojsurT 29 pimH (7961) Trega (6£61) Aaysury (€C6L) IIUIYNIN (TEGI) UTeMgoeyy 29 Aafsur] (Z961) [PUIN : (TS6L) Aepsury 29 pimH ‘(QE461) SUIT ‘pae 39 (LZ61) 3JOYSIY (86661) PUNJAT ‘pone 39 (LZ6T) JJOYSIE paysıygndun (FS6T) Hoyspaors (7961) PUNJAT (96561) PPunJarT uoss31J sadixayy (WI) “VY UOSSIIT FIND (esoydoyuessıw) vzoydopuy Avg vidnaqv viogdoqiupy uossaig vurgin ‘Y CID) vuvipsofurys ‘y ayepoquny rdagsun "y “TPIO euerp -JOJUEIS "Y ‘UOSSIIZ TISPIEMPS "Y ‘dds zayzo snorzea 29 Avg vidnige eroydoyuy (issoy) 2/44 vypsaump (day) vasvjora you (day) euoSrqre ejpdauy Crus) rruquif ijd py “JOY stjearog "YW © (‘zueg) wernseunrpenb exogdomuy SELIN Aer (smoq) pwuozoauumg[ you (ID) sesuonang vysaup jnpqnop vonisod ua) (7 (uossaJ) wejndeu “WI peowysy 122450307 (uossa15)) vidnasaqur (vgdsowojsapayy ) (uossa1) vue (eydıowopapp) ASSUTT 2722704042 X (25194) suwsuna ‘Wary SUI UNS (day) snsowea “gar surgsarıd (day) smeqzo CTD) sus AR (CPE) ssuouroquer avipurjjogaraou 260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 other cases the associations recorded are merely suspected. Nevertheless, a survey of the literature reveals that a substantial number of records are well founded and based on a study of the biology of both parasite and host, the assessments of true parasitism having been accomplished by rearings or otherwise. Consequently, in the Table all names of well-established associations have been printed in bold type. DISTRIBUTION OF REGIONAL GENERA AND SPECIES Leaving the widely spread Thyreus out of account, and with the exclusion also of the large genus Melecta — which is abundantly represented in the Mediterranean basin and West Asia — all Old World members of the tribe occurring in temperate climatic regions are rare in collections, despite their striking body pattern. The locality records are few in number and scattered over wide areas, while hardly anything is known of their habita- tions. This applies not only to Paracrocisa and Pseudomelecta, but also to the isolated Ethiopian genera Afromelecta and Acanthomelecta, two taxa founded on solitary species represented by few individuals only. We are somewhat better informed geographically about the occurrence of Eupavlovskia and Protomelissa, whose ranges, as far as at present known, are roughly outlined in the accompanying maps, in conjunction with the dis- tribution of their hosts, i.e. the anthophorine genera Habropoda and Elaphropoda (see maps I and II on p. 268 and 271, respectively). For a more detailed account of Eupavlovskia, see Lieftinck (1969). It will be seen that Protomelissa is mainly tropical in distribution. Its species exhibit a colour design that deviates much from that of the others, in such a way, in fact, as to attract little attention in the field. With few excep- tions, the remainder are characterized by a pubescent pattern of black and snowy-white, many of them when freshly emerged being extremely handsome and conspicuous bees. KEY TO THE MELECTINE GENERA OF THE EASTERN HEMISPHERE 1. Marginal cell of fore wing usually markedly shorter than three submarginal cells combined, at most equal in length to, usually much shorter than, the distance separating its apex from wing tip, whether or not exceeding third submarginal cell. Body black-haired or spotted with white and/or blue, pubescence never pale yellow, orangish, fulvous or dark brown. . . 2 — . Marginal cell long, usually little shorter aa tien submarginal CE omai longer than the distance separating its apex from wing tip, and exceeding third submarginal cell (Fig. 11). Body lacking well defined white or blue pubescent spots: whole dorsal surface and sides of thoracic segments clothed densely with long erect pubescence entirely concealing surface. First gastral tergite markedly shorter than second, lacking tufts of longish erect hairs; abdominal pubescence short, generally dense, appressed and feathery, but never forming spots and usually not much different in colour from that of underlying surface. Head only one-third as deep as its breadth across eyes, posterior angles not developed. Face rather short, slightly to moderately protuberant, i.e., height at apex of clypeus in side view about one-third to four-sevenths greatest diameter of eye at the same level; clypeus broader than long, convex dorsally. Labrum broader than long, often emarginate anteriorly. Mandibles curved, shorter than length of eye, inner margin unidentate or with angular projection far beyond halfway length. Inner orbits nearly straight, sub- M. A. LIEFTINCK : Old World Melectine bees 261 parallel or slightly converging anteriorly. Maxillary palpi 6-segmented, long and slender, subequal in length to antennal scape. Antennae rather long and slender, often exceeding tegulae, flagellar segments variable, but in male up to almost three times as long as broad; rhinaria absent. Mesoscutellum short and convex, much broader than long, with distinct middorsal carina; dorsal lobes armed with a robust erect and curved spine, as long as or shorter than surrounding pubescence and directed caudad. Parascutella small, subtriangular, convex. Wings lightly infuscated. Legs rather thin and slender, of simple structure; hind tibial spurs almost straight, finely serrulate, outer spur only little shorter than inner. Tarsal arolia well devel- oped, though occasionally of minute size. Male terminalia, see specific descriptions and figures. Pygidial area of female broad, subtriangular and plate-shaped, not covering entire dorsum of tergite. Body size small to medium, 10.5—13.5 mm. Hab.: Himalayan region to Taiwan and Java. . . Si: Protomelissa First gastral tergite usually distinctly longer than er in normally exposed position, or at least equal in length to the latter. Tarsal arolia vestigial or absent 3 . First gastral tergite markedly shorter than second. Tarsal arolia conspicuously present, claws usually divergent. Maxillary palpi distinct, 5—6 segmented, as long as, or little shorter than, three La segments of labial palpi combined, exceeding 1 mm in length . . 5 Scutellum plate- ike: flat or A so, die Lia with dorsal Bre abruptly. and acute-angularly divided from forward-slanted or incurved posterior surface, never armed with a pair of cylindrical or abruptly raised spines or tubercles; more rarely slightly convex or bituberculate and, though not markedly projecting beyond meta- notum at middle, yet distinctly angulate in profile, with the prominent lateral angles directed caudad and overhanging metanotum laterally. Parascutella triangular, on a level with scutellum or almost so, occasionally convex dorsally. Tarsal arolia absent or replaced by a vestigial orbicula. Face protuberant, height at apex of clypeus at least equal in profile to greatest diameter of eye at the same level. Maxillary palpi wanting or rudimentary (1-2 segmented) and only occasionally made up of more (3-4) short segments, whole palpus shorter than, or about as long as, distal two segments of labial palpus combined and not exceeding 1 mm. Mandibles with inner margin strongly unidentate about halfway length. Antenna normal, thickness and length of flagellar segments variable but never unusually attenuated; rhinaria often present in both sexes and rarely wanting in male. Wing membrane often obscurely spotted and frequently very dark. Outer face of mid tibia often somewhat flattened and/or broadened, and nearly always with dense pad of short felt-like pubescence. Integument of thorax above at least partly exposed; dorsal pubescence short and frequently dense, but integument never completely hidden from view. Abdomen with short, sparse black hairs variegated with pubescent spots composed of longer, but predominantly appressed, branched, feathery or scale-like hairs, either pure white or coloured various shades of blue and/or green. Size very variable. Hab.: Old World subtropics and tropics, with radiations into the southern Palaearctic ee . Thyreus : Scntellum maaike) but either flattened nor plate-like, SE segment about twice as broad as long, strongly but gradually declivous and impressed posteriorly; dorsal surface not overhanging any part of metanotum, rounding into vertical posterior surface, the swollen parts on either side of median concavity armed with compressed 262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 tooth or spine placed far laterad and directed caudad; parascutella inconspicuous, convex, usually hidden from view by dense appressed pubescence. A conspicuous patch of white appressed tomentum at middle of scutellum posteriorly. Tarsal arolia rudimentary. Face broad. Labrum subquadrate, anterior border bituberculate with distinct crescentic emargination; surface partly or entirely clothed with dense brush of stiff hairs. Clypeus convex, little prominent, much shorter in profile than greatest diameter of eye (Fig. 46). Eyes converging anterad, inner orbital margin straight and upper portion not or scarcely incurved; in side view widest below middle of their length, much more broadly rounded anteriorly than above. Mandibles strongly curved, inner margin edentate or with vestigial tubercle placed about midway or slightly in advance of their length. Malar space linear. Antennae slender, un- modified, similar in both sexes, barely attaining tegulae (female), or slightly longer and reaching to beyond middle of latter (male); scape curved, with short appressed pubescence; flagellar segments normal, but longer than broad, rhinaria wanting. Legs normal, shaped similarly in both sexes, mid and hind basitarsi of male un- modified, covered externally with short, dense, appressed white pubescence; hind tibial spurs relatively short, gently curved, inner spur not much longer than outer. Fore wing membrane partly obscured; marginal cell elliptical, rather attenuated distad, only slightly shorter than distance from apex to wing tip, exceeding third submarginal cell. Front of head thickly white-haired, pile on clypeus decumbent but less dense and very short anteriorly; thorax variegated with semi-erect black and white pubescence only partly hiding surface on mesonotum and scutellar area. Abdomen dull, especially on dorsum, surface finely closely punctate, but punctation invisible where covered by dense, appressed, finely branched hairs forming well- defined white spots. Gastral tergite 7 of male rounded apically. Exposed surfaces of male gastral sternites 2-6 uniformly pubescent; no indication of a conspicuous subapical comb of thick brushy hairs projecting from beneath sternal plates 3-5. Sternite 7 of male constricted basally, distal portion plate-shaped, more or less bilobate and with dense frings of strong bristles; apex of 8 subtruncated and fringed with longish hairs (Fig. 51, 56). Genital capsule of large size and complex structure (see specific descriptions). Pygidial plate of female triangular, with broadly rounded apex EDEN EIA CREER IA OENE Maxillary palpus slender, 6-segmented, 2nd to 4th longest (Fig. 55). First gastral segment distinctly longer than second in normally exposed position (male), or second segment, when extended, almost equal in length to first (female). Convex scutellar lobes armed on either side with an enormous, slightly downcurved, laterally compressed spine, which is directed straight back, equals middorsal length of scutellum, and bears a strong, longitudinal dorsolateral carina. Tarsal arolia con- tracted to a plumose or bristly rudiment. Hab.: East Africa (Tanzania). . . . . Acanthomelecta . Maxillary palpus rudimentary, consisting of 1-2 minute segments, which together are equal in length to, or shorter than, the two apical segments of labial palpus combined. First gastral segment a little shorter than, or subequal in length to, the second. Convex scutellar lobes armed with a pair of slightly downcurved and laterally compressed teeth or spines, which are slightly divergent and at most half as long as midlength of scutellum; dorsolateral margin of each strongly longitudinally carinate. Tarsal arolia wanting, replaced dorsally by an orbicular fan of long and 6. M. A. LIEFTINCK : Old World Melectine bees 263 strong bristles, and eee i a fringe of bristles suggesting a planta. Hab.: South and East Africa . . . „u. = ue Afromelecta Soft raised pubescence covering oral ee a exceptionally long and dense, especially at sides, the thick velvet-like pile composed of feathery hairs con- cealing all of the surface inclusive of posterolateral pronotal tubercles and scutellar spines. Basal propodeal triangle glabrous, except few short longitudinal striae laterally at base. Fore wing greatly expanded towards apex, distal margin strongly convex, tip blunt; marginal cell short, almost regularly elliptical, over three times as long as broad, not or only little surpassing third submarginal cell. Body compact, thorax large and globular; abdomen even more abruptly pointed than in Melecta and Pseudomelecta. Maxillary palpus 6-segmented. Mandible unidentate, tooth rather large and placed near apex in male, rather smaller and blunt, situated slightly beyond halfway length in female. Antenna long and slender in both sexes, scape shortly pubescent, segment 4 always longer than 3 and more than twice as long as broad, next flagellar segments also attenuated, about twice as long as broad, the distalia frequently slightly crenulated in male; rhinaria wanting. Thoracic dorsum closely punctate, scutellum with a pair of robust, straight, upturned divergent spines hidden by surrounding pubescence. Legs dull, outer faces of all parts finely reticulate- punctate, conspicuously lacking spines or bristles, clothed densely with short, pre- dominantly black pubescence almost concealing surface, only femora below fringed with long hairs. Mid tibia of male not expanded, lacking conspicuous external pad of short felt-like pubescence; hind tibia of male expanded and swollen distad, with strongly produced, robust antero-apical process; hind basitarsus of male modified, laterally compressed, rather abruptly expanded and hollowed out externally, that of the female subparallel-sided. Hind tibial spurs long, innermost spur undulated and much longer than outer, which is straight, both microscopically serrulate. Inner rami of mid and hind tarsal claws slender, neither flattened nor plate-like. Gastral tergites finely superficially tessellate-punctate, surface dull or slightly shining. Pubescent pattern variable: body either entirely jet-black, or thorax with white or greyish- yellow anterior mesothoracic collar, long black and white lateral tufts behind wings, lateral tufts of long raised hairs at extreme base of tergite 1, and snowy-white appressed pubescent spots on each side of succeeding segments. Pile on mesonotum never forming grey-white and black spots or streaks. Tergite 7 of male subtruncated, apex clothed with appressed tomentum. Sternite 7 very slender, with narrow, widely divergent arms and bilobate apex, the lobes fringed with strong bristles; sternite 8 with well developed postmedian ridges, apex with tufts of long feathery hairs. Hab.: Western Mediterranean to Caucasus . . . . . + . Eupavlovskia . Raised pubescence covering dorsal and lateral a segments shorter, often dense and feathery but only very rarely completely concealing surface, the integument remaining visible at least on middorsum and frequently also on parts of the sides; black and white pubescent pattern variable. Body shape and size, and form of wings very variable, but combined structural characters of antennae, legs, and male apical sternal plates not as described above for Eupavlovskia . . . . Pet 6 At least posterior portion of mesonotum and entire scutellum integument well visible under scanty pilosity. Mesonotum anteriorly, mesopleurae above, a sub- circular area in front of parascutella at each side of mesonotum, and dorsolateral metanotal area, clothed with longer, but mostly decumbent, silky hairs frequently 264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 forming condensed patches of soft, silvery-white pubescence. Abdomen variable, tergites usually dull and almost lustreless, clothed with minute dark hairs and often variegated with pure white pubescence forming spots or definite patches of dense, appressed silky hairs of somewhat greater length; tergite 1 never with tufts of long raised hair on each side at extreme base (except Pseudomelecta chalybeia). Ab- domen occasionally more shining and almost bare (some species of Pseudomelecta). Sternites 2-6 (or at least 3-5) of male with subapical fringe of strong, backward directed, black bristles, which often form a dense brush. Legs with short, generally scanty, pubescence, except longer fringe at fore femora posteriorly and, more sparsely, often also at basitarsi. Inner rami of mid and hind tarsal claws always much shorter and broader than outer, the former usually flattened and often axe-shaped or squarely cut off. Scutellum broader than long, raised and greatly swollen, well marked off anteriorly from mesonotum and convex parascutella by deep furrows; dorsal surface (inclusive of well developed backward directed processes) on a level with mesonotum in side view, strongly declivous posteriorly, but whole structure not or only little overhanging posterior thoracic segments. Antennal scape naked or covered with minute appressed pubescence, rarely with sparse fringe of long erect hairs. Mandible with single, robust, interior tooth beyond halfway length. Malar space linear or wanting. See also under couplet 6b, of Melecta. . . . . 7 . Raised pubescence on thorax rather long and dense, usually covering all parts of dorsum and sides: although integument remains visible in places, most areas are hidden from view, including the scutellar spines, which are frequently raised, but often vestigial and only rarely as long as the surrounding pubescence. Pile entirely black or, more often, predominantly grey-white (rarely brownish-white) on mes- onotum anteriorly, becoming black and/or white posteriorly (male), or forming alternating grey-white and black blotches and streaks on anterior part of dorsum (female). Gaster moderately to strongly shining, frequently almost polished, disk of tergites finely, sparsely punctate and with short scanty pubescence; tergite 1 in most species with tufts of long erect white hair laterally at extreme base, sides of this and/or following tergites in addition spotted with condensed patches of ap- pressed white pubescence. Sternites often with long, strong subapical hairs, but 2-6 (or 3-5) of male lacking dense combs of thick stiff bristles pointing caudad. Anten- nae normal, relative lengths of segments varying among species; scape frequently fringed with long hairs; flagellar segments (except two basal ones) never distinctly attenuated or much longer than broad, often thick and distinctly transverse, especial- ly in male; rhinaria almost invariably present in male. Legs variable, but hind tibia of male unmodified; outer faces of tibiae and tarsi often tuberculate and/or denticulate; mid tibia of male frequently expanded and with dense external flattened pad of short felt-like, white or black, pubescence; inner hind tibial spur not undulated; hind basitarsus of male not modified, but outer face frequently concave and apex occasionally distinctly produced externally. Shape and armature of scutellum, as well as of tarsal claws, variable. Numerous species of very variable appearance and size. Hab.: Old World forms widely-spread in temperate regions. . . . Melecta N.B. — In a number of aberrant Melecta, the integument of the thoracic segments is well visible under much shorter and more scanty pubescence, while the gastral tergites are lustreless, finely punctate and clothed densely with minute hairs, the extreme base of tergite 1 at the same time lacking the characteristic lateral tufts of long raised hair. Members of this species M. A. LIEFTINCK: Old World Melectine bees 265 group are extremely similar superficially to Paracrocisa, the shape of the scutellum also being almost identical. Despite this similitude, they have the following negative characters in common by which they can be distinguished from Paracrocisa. (1) labrum in both sexes not markedly broader than long, usually distinctly longer than broad; (2) antennae of normal length, scape in male with lateral fringes of long erect hairs, the flagellar segments not unusually attenuated and with distinct, impressed rhinaria; (3) inner rami of mid and hind tarsal claws of male and female never distinctly plate-like, axe-shaped or squarely cut off; (4) gastral sternites 3-5 of male without dense subapical combs of thick, suberect bristles pointing caudad; (5) hind tibia of male not markedly expanded, apex not hollowed out and toothed; and (6) apical sternal plates and genital organs of male not at all shaped and modified as described and figured for Paracrocisa. The females are less easily recognised, those of Melecta usually having a more protuberant face, a longer labrum, and the exposed portions of abdominal sternites more broadly impunctate. Body small (length 8-12.5 mm), compact; abdomen thick-set, ovoid (e.g., length: breadth ratio 10 : 7.5-8.6). Marginal cell of fore wing short, elliptical, about 2.2-3 times as long as broad, apex broadly rounded or subtruncated; sides of second submarginal frequently closely approximated above but only rarely fused together or stalked at anal side of marginal; third submarginal distinctly smaller than first and generally a little higher than long. Antenna moderately long, usually reaching to slightly beyond anterior margin of tegulae; shape varying between species: either thick and strong, with flagellar segments distinctly transverse and bearing deeply impressed, crescentic or horseshoe-shaped rhinaria (male), or thinner with the dis- talia longer than broad (female); or else, much slenderer and almost alike in both sexes, the flagellar segments longer than broad, and with male rhinaria only feebly indicated. Legs short and robust, femora and tibiae of male slightly more expanded than in female, but apices of male tibiae and basitarsi lacking any specializations; mid basitarsi of male invariably more or less expanded and flattened, with external pad of dense appressed pubescence. Hind tibial spurs more or less curved, not wavy, inner spur not considerably longer than outer. Maxillary palpus 6-segmented, seg- ments varying in length, the two terminalia occasionally fused together, 2 and 3 usually longest. Labrum very small, subquadrangular, at most half as broad as lower interorbital distance. Eyes broader and more evenly rounded above than in Para- crocisa, a little shorter than upper interorbital distance; inner orbits straight and converging anterad. Texture of abdomen variable, tergites moderately shining and punctate, but often lustreless, with very closely set fine punctures; premarginal (sub- apical) brush covering sternites 3-6 of male composed of evenly distributed, stiff suberect bristles, fewer in number and less conspicuous than in Paracrocisa, hairs not upcurved apically. Sternite 7 of male with strongly protuberant lateral angles at end of divergent arms; disk broad and flat, not at all constricted basally, sub- rectangular or more or less tapering posteriorly, lateral margins not thickened and only weakly sclerotised, apex beset with few strong bristles. Sternite 8 with poorly marked and strengthened subapical angles at base of slightly prominent apex, the hind margin of which is undulated, carrying few bristle-like hairs on either side (Fig. 20, 24, 26). Genital capsule compact, small and spherical, gonocoxites broad- ening gradually posteriorly, with rather short, strongly converging gonostyli; no membranous flap at base of gonostylus; dorsobasal and ventrobasal processes var- iable, conspicuous, but not strongly modified, densely pubescent and bristled (Fig. 21, 25, 27). Hab.: E. Turkey (Armenia) eastward to Mongolia. Pseudomelecta 266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 — . Body size moderate (length 12-17 mm), elongate; abdomen slender, less abruptly pointed than in Psewdomelecta (e.g., length : breadth ratio 10 : 6.6). Marginal cell of fore wing narrower, about 3.3 - 4 times as long as broad, with bluntly pointed apex; third submarginal cell frequently almost equal in size to first. Antenna long and very slender, though little surpassing tegulae (male), or shorter and reaching anterior margin of same (female); segments 3-5 slightly more than 114 times as long as broad and of about the same length, remaining segments only little shorter, all smooth and rather shining below, with shallow longitudinal impressions, but lacking distinct rhinaria (male); or shorter, with separate flagellar segments about 11/3 times as long as broad (female). Legs strong, hinder pair modified in male, normal in female. Hind tibia of male considerably expanded and swollen towards apex, which is hollowed out below implantation of basitarsus, ending in a variously shaped ventral process carrying articulated spines; spurs originating from the inside just above the truncated apical border, the inner spur distinctly wavy, much longer than outer; hind basitarsus strongly compressed laterally and a little outbent, its external surface distinctly hollowed out. Maxillary palpus 6-segmented, long and slender, about as long as last three segments of labial palpus combined, 1 only half as long as next segments, which are subequal in length, 6 only little shorter than the preceding. Labrum larger than in Psexdomelecta, much shorter than its width at base and broader than half the lower interorbital distance. Eyes very large, as long as, or a little longer than, upper interorbital distance, elongate-oval, broadest slightly in advance of their middle and narrowest above; inner orbits converging anterad, straight or slightly convex inward. Abdomen generally dull, evenly finely punctate from base almost to apex of tergites; premarginal (subapical) brush covering sternites 2-6 of male conspicuous, composed of thick bristles, the apical ones comb-like and usually with upturned tips. Apex of tergite 7 of male sub- truncated or bluntly rounded. Sternite 7 lacking prominent lateral angles at end of widely divergent arms, abruptly and very strongly constricted at base of disk, which is at first long, narrow, and subparallel-sided, then broadens rather abruptly to form a plate-like apical lobe bearing numerous strong, closely set, radiating bristles on either side. Sternite 8 subtriangular, surface basally with well marked ribs and prominent, heavily sclerotised subapical angles at base of protuberant and equally strengthened apex, the hind margin of which is finely setiferous (Fig. 40, 42, 44). Genital capsule oval and of large size, gonocoxite abruptly broadened posteriorly, with large membranous flap at base of long gonostyli, and intricately shaped basal processes carrying a variety of bristles and fine hairs (Fig. 41, 43, 45). Pygidial plate of female slender, narrowly triangular, about 21/, times as long as its width at base, apex rounded. Hab.: N.W. Africa to E. Kazakhstan . Paracrocisa Discussion OF GENERA AND SPECIES (EXCLUSIVE OF Melecta Latr., 1802) Thyreus Panzer, 1806 This is probably the most advanced and diversified genus of the tribe, the members of Thyreus showing a number of specializations and reductions not found elsewhere. The relatively great length of the first gastral segment as well as the obliteration of the maxillary palpus and lack of arolia, are presumably all characters indicating the highest M. A. LIEFTINCK: Old World Melectine bees 267 stage in the evolution of the Melectini. With the exception of the numerous species peculiar to the Ethiopian fauna — which are still in great need of further study — most Thyreus occurring in the rest of the world were reviewed in a series of articles published by Lieftinck (1959—1968). All Anthophorini whose association with Thyreus has been either ascertained or may reasonably be expected, are listed in the Table on p. 258 of this paper. Type species: Nomada scutellaris Fabricius, 1781. Eupavlovskia Popov, 1955 A small genus that has recently been treated in full (Lieftinck, 1969). The silky pubescent pattern of Eupavlovskia strongly recalls that of certain aberrant Eurasian members of the polymorphic and widely spread genus Melecta. Yet the genus differs fundamentally from all species of Melecta in several structural characters suggesting a closer affinity with Paracrocisa than with any other tribal genus. There are only two species, which are apparently confined to southern Europe and parts of West Asia, the centre of their distribution lying in the peninsular countries of the north-east Mediterranean (see Lieftinck, loc. cit., fig. 8, map). Eupavlovskia is parasitic on the larger-sized species of Habropoda, i.e, the more typical members of that genus. The approximate range of both units has been roughly outlined in Map I on page 268 of this paper. Further towards the east, these anthophorines are replaced by a far greater number of somewhat smaller ‘Habropoda' showing characters approaching those of the Nearctic Emphoropsis. In point of fact certain Eurasian mem- bers of this group are rather intermediate morphologically and are suspected to be the hosts of Protomelissa, discussed hereafter. On a collecting trip to Rodos, in April 1971, I took some E. obscura simulatrix Lieft. in company with Habropoda tarsata (Spin.), the latter being common all over the island. On the nearby island of Kos, opposite the Turkish coast, the same Expavlovskia occurred together with both H. tarsata and zonatula, but of the last only a single male was observed. Unfortunately no nesting sites were found of either of the host bees. Type species: Melecta funeraria Smith, 1854. New localities, in addition to those enumerated in my former paper, are: E. funeraria (Smith). — Italy: @, Calabria, Antonimina, 1905 (MNB). Greece: 9, Naxos I. (MNB). E. o. obscura (Friese). — Italy: 9, Naples (as M. aterrima Lep.) (MNB); S 9, Apulia, Sporgano, IV-V. 1933 (MNB). E. o. simulatrix Lieft. — Greece: J 2, Corfu (as M. funeraria Smith, det. Alfken) (MNB). Protomelissa Friese, 1914 Protomelissa Friese, 1914, Dtsch. Ent. Z. : 322—323 (two species: P. sauteri (Fr.) & 2 Formosa, and P. iridescens Fr, & Formosa). — Cockerell, 1929, Ann. Mag. Nat. Hist. (10) 4 : 133 (note, not seen). — Sandhouse, 1943, Proc. U.S. Nat. Mus. 92 : 592 (not seen; designation of type species P. iridescens Friese, 1914). — Lieftinck, 1944, Treubia, hors sér.: 59 (note, not seen). — Michener, 1944, Bull. Amer. Mus. Nat. Hist. 82 : 287 (note, not seen). — Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21 : 322, 333 (notes, not seen). Callomelecta Cockerell, 1926, Ann. Mag. Nat. Hist. (9) 18 : 621—622 (2 Malay Peninsula; TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 268 vpodosqdviq pue rpodosqry e1aua3 }soy ours -oydoyjue zo aduer [ensed uo pasodwunodns (aed ur 9YSTI) 155//2440/04] pue (aur, Arepunog Jo 3791) wysaojardug eruo] aum>ajaw Jo uonnqinsiQ ‘I dem BSSTTSWOYOId erysnoInedn3 epodoıyde13 2 epodorqey M. A. LIEFTINCK : Old World Melectine bees 269 type species by monotypy C. pendleburyi Ckll., 1926). — Sandhouse, 1943, Proc. U.S. Nat. Mus. 92 : 533 (not seen; type-species C. pendleburyi Ckll., 1926). — Lieftinck, 1944, Treubia, hors ser. : 58—62, fig. & pl. (full generic & specific descr. & 9, type-species C. pendleburyi Ckll). — Michener, 1944, Bull. Amer. Mus. Nat. Hist. 82 : 287 (note, not seen). — Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21 : 333 (note, not seen). The first described species in this genus is Anthophora sauteri Friese, 1911, originally known from a single male “von Taiwan (Formosa), Sauter leg.” When Friese (1914) had received examples of both sexes, he proposed the new generic name Protomelissa, to hold it. The generic diagnosis, brief and insignificant as it is, starts as follows: “Eine neue Schmarotzer-Bienengattung, die noch vollkommen den Anthophora-artigen Habitus bewahrt hat und ein schönes Mittelding zwischen dieser Gattung und den typischen Melecta- und Melissa-Arten darstellt. Protomelissa sauteri wurde von mir 1911 nach einem defekten & als Anthophora s. beschrieben, nach Bekanntwerden des 9 treten aber alle Eigenschaften der Schmarotzerbiene zweifellos hervor und lassen sie als ein vortreffliches Bindeglied zwischen Anthophora und Melecta-Melissa erscheinen.” (loc. cit.: 323). In the next pages Friese describes the female and adds a few characters of the male, starting his description of P. sauteri again with the following sentences: “Diese von mir nach einem ¢ als Anthophora beschriebene Art gehört nach dem Bekanntwerden des 9 zu den Schmarotzerbienen, sie liefert ein gutes Bindeglied zwischen Anthophora und Melecta. Der Habitus ist ganz Anthophora-artig, auch die Behaarung, doch fehlt im Q der Beinsammelapparat (= Scopa). Ich gebe hier eine neue Beschreibung, zu- gleich die Diagnose für das 4 erweiternd.” The type of A. sauteri is not “defekt”, as stated by Friese, but a specimen in good condition (see below). However, it may have been the reason why Sandhouse (1943), instead of selecting the much better known P. sauteri, designated P. iridescens Friese as the type species of Protomelissa. This type designation is incongruous with the facts that (1), P. sazteri is not only the earliest but also the fullest described species of which both male and female were known, the diagnosis of Protomelissa, as stated explicitly by Friese himself, being based primarily on characters of the female; and (2), because the unique type male of P. iridescens does not quite fit the generic diagnosis and has apparently become lost, the description of it being even more fragmentary than that of P. sauteri. See also p. 277 under P. iridescens. It has now been found that P. sauteri is conspecific with Melecta formosana Cockerell, 1911, described about two months earlier than sauteri, so that the former name should replace it. At the time of redefining Callomelecta Ckil. (Lieftinck, 1944), no species of Proto- melissa were known to me, but in 1970, while studying three of its members in the Berlin museum, it soon became clear that the two genera are inseparable. A re-char- acterization of Callomelecta was given by me in 1944, and this elaborate diagnosis applies equally well to Protomelissa in almost every respect. In general facies all species presently included in it are strikingly alike, while they agree also in the structure of the mouth- parts and venation; lastly, the male terminalia and female pygidial plates of Protomelissa correspond closely with those of Malaysian Callomelecta. The differences are mainly found in the shape and pubescence of the antennae and tarsal claws, which seem to be of specific rather than generic importance. As may appear from the photographs (Pl. 1 Fig. 1-2), the members of Protomelissa are entirely unlike any of the Old World melectines, differing from all in the unspotted body and vividly coloured pubescence, features suggestive of their hosts and leading to 270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 the opinion that they have preserved some primitive features peculiar to the ancestral anthophorini. It is tempting to speculate upon the possible relation between parasite and host throughout their range. There seems to be little doubt that the equatorial members of this Asiatic genus (i.e., Callomelecta olim), are restricted to the submontane and montane forest zones of Malaysia, and that they are special parasites of the anthophorine Elaphropoda. Three species of the latter have been found associated with them in such different localities as the Malay Peninsula, Sumatra, and Java. P. habropodae (Ckll.), on the other hand, was captured by its describer in the mountains of Thailand simul- taneously with Habropoda sutepensis Ckll. Nothing, however, is known with certainty of the more northern Protomelissae, described from the Himalayan range and Taiwan. There are about four nearly interrelated species of Elaphropoda and at least five or six Habropoda (excluding a few undescribed ones), which are distributed throughout the eastern Himalayan mountains. From the corresponding locality records and dates of capture it is obvious that some members of either genus inhabit the same area and may even occur side by side (see maps). In an easterly direction, the two anthophorine genera are each of them represented by at least two species in the mountain provinces of East China (Fukien); but whereas Habropoda tainanicola Strand, H. bucconis (Friese) and a third — possibly undescribed — species have succeeded to reach Taiwan — in which island they occur at much lower levels than on the mainland — no single species of Elaphropoda has turned up yet in Taiwan. Their absence is significant because, as we will see, no less than three Protomelissae do occur in the island and strongly suggest their association with the two Habropodae just mentioned. The absence of Protomelissa on the mainland of China and the Indochinese provinces does not, of course, preclude the possibility of their occurrence in the intervening countries as it can easily be explained by their rarity and also the insufficient exploration of the Chinese mountains. This is also the opinion of Dr. T. C. Maa, who recently told me in a letter that the lacunae in his very rich Fukien collections of bees are, perhaps, due to the fact that collecting was done in all seasons for seven consecutive years, but only around a field station set up at 1600 metres; only occasionally he and his assistants visited the lowland as well, thus missing the right season and localities for Protomelissa. KEY TO THE SPECIES OF Protomelissa (The females of P. habropodae and iridescens, and the male of tricolor, are unknown) 1. ‘‘Integument of abdomen shining bright clear ferruginous, clothed above with short, appressed, pale fulvous tomentum on apices of second and following segments. Thorax with abundant, long, erect fulvous hair. Clypeus black, but densely covered with appressed pale hair. Flagellum of antenna not very long, barely reaching tegulae, apical joint curved. Female unknown.” (transcribed from Cockerell). Hab.: Thailand (Siam) 1 tt: habropodae — . Integument of abdomen at least ln basal a di He two segments dark brown or black and clothed above with appressed, light brown to brownish black or black tomentum, the hairs varying in length: either short and simple, or longer and finely branched; integument of distal portion of tergites often more or less ferruginous, and if so, then distal portion of antennal scape clothed in front with long, raised, pales pubescence” Hate ta n> sen Re) ee EN AE Me M. A. LIEFTINCK: Old World Melectine bees 271 HABROPODA & ELAPHROPODA PROTOMELISSA Map II. Distribution pattern of the melectine genus Protomelissa superimposed on the eastern range of the suspected anthophorine host genera Habropoda and Elaphropoda 272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Dense pubescence covering thoracic segments above unicoloured brownish- to orange-yellow, a or xanthine-orange, composed of long, finely branched raised hairs . . EP 3 . Dense pubescence ee onde men above picelowed! and composed of still longer, finely branched, raised hairs: either pale yellow with a transverse blackish brown median pubescent band between the wings, or else, dark brown anteriorly as far back as base of scutellum, and canary-yellow posteriorly. . . . . 6 “Integument of abdominal segments above and underneath red-brown, the bases black; tergites finely punctate, dullish, with slight metallic reflex; pubescence short, reddish, felt-like; apex of tergite 7 of male straight cut off. Labrum squarish, anterior margin shallowly emarginate. Antenna red, second flagellar segment (3rd antennal) much shorter than third (4th antennal), all flagellar segments of simple structure. Head and thorax above clothed densely with long reddish yellow (“rot- gelb’) pubescence. Legs blackish brown. Female unknown.” (transl. from Friese). Hab.: Taiwan (Formosa) . . . bis 0 iridescens . Combined characters not as above. Inner ramus er mid xs hind tarsal claws short, thin, and almost square; tarsal arolia conspicuous. . . AR: 4 Pubescence on dorsum of abdomen short and sparse, ara dade ite not plumose and not concealing surface, contrasting strongly in colour with long, feathery, bright orange-red pubescence on head and thorax. Antennal segment 3 a little shorter than 4 (male), or more markedly so (female). . . . 5 . Pubescence on dorsum of abdomen short, but much denser! a light ore, forming more or less distinct colour bands of appressed plumose hairs partly hiding the surface and not much different in colour from the long, feathery pubescence covering head and thorax. Labrum emarginate anteriorly. Apex of tergite 7 of male bituberculate, the approximated lobes separated by a crescentic emargination. An- tennal segment 3 subequal in length to 4 USE or a little shorter than this (female). Hab.: W. Java. . . . . insidiosa Anterior margin of labrum almost straft or an liphilye concave. Integument of body and legs reddish- to brownish black. Dense feathery pubescence covering dorsum of thorax capucine-yellow to mikado-orange. Apex of tergite 7 of male produced in two more or less triangular lobes, which are irregularly truncated and separated from each other by a deep U-shaped emargination, which is about equal in size to each of the projecting tubercles. First gastral tergite of female clothed sparsely with short, light ochraceous-buff tomentum, following tergites with very short brownish black hair, 2—4 moreover with transverse lateral patches of light ochraceous-buff tomentum. Hab.: Malay Peninsula. . . . . pendleburyi . Anterior margin of labrum distinctly emarginate. Integument of body and legs black. Dense feathery pubescence covering dorsum of thorax bright xanthine-orange. Apex of tergite 7 of male produced in two more or less triangular protuberances separated from each other by a shallow emargination. First gastral tergite of female clothed with short, appressed, ochraceous-orange tomentum forming a narrow, trans- verse and obliterated band, following tergites clothed sparsely and evenly with brownish black hair. Hab.: Sumatra. . . chive vulpecula Long pubescence covering head and dorsum of pro- and mSsönohım dark chocolate- brown as far back as just behind tegulae, all the rest of thorax clothed densely with yellow hairs. Labrum very short, almost twice as broad as long and widest at middle, M. A. LIEFTINCK: Old World Melectine bees 273 anterior margin shallowly emarginate (Fig. 4). Front of antennal scape pubescent, but lacking tuft of much longer raised hairs at distal portion; segments 3 and 4 much longer than broad, 3 only little shorter than 4. Inner ramus of mid and hind tarsal claws short, plate-like, axe-shaped and almost square; tarsal arolia vestigial. Male unknown. Hab.: Assam. . . . . tricolor —. Long pubescence covering head and ye mainly on. in mesonotum with broad, transverse, blackish brown pubescent stripe between the wings. Labrum much longer and only little shorter than its greatest breadth at base, anterior margin deeply emarginate (Fig. 1, 3). Front of antennal scape clothed densely with pale raised hairs, which in the male form a conspicuous compact brush; segment 3 very short, only about one-third as long as 4 (male), or a little less than half as long as this (female); flagellar segments distinctly crenulated posteriorly (male, Fig. 2), or normal (female). Inner ramus of mid and hind tarsal claws longer, laterally com- pressed, but claw-like, not squarish; tarsal arolia conspicuous. Scutellar processes glossy black, in the form of long, very robust, slightly divergent, curved thorns, which are occasionally more or less soother-shaped, with adruptly pointed tips . 7 7. Scutellar spines in dorsal view entirely surrounded and hidden from view by very long and dense pale pubescence; no small tuft of long dark hairs just in front of the spines and no dark-tipped hairs just beside tegulae. Labrum a trifle longer and less deeply excised anteriorly (Fig. 3). Apical sternal plates and genitalia of male as in oe 7—8. Hab.: from Kumaon (N. India) as far as Shillong (Assam) tee i himalayana —. tell es in dorsal view not dui nee the black tips remaining partly visible between very long, but slightly less dense, pubescence, which is pale-coloured, save for a small, isolated tuft of equally long dark hairs just in front of the roots of the spines; also a few dark-tipped hairs just beside tegulae. Labrum a trifle shorter and more deeply excised anteriorly (Fig. 1). Apical sternal plates and genitalia of male as in Fig. 5-6. Hab.: Taiwan (Formosa). . . . formosana Protomelissa himalayana (Bingham), comb. nov. (Fig. 3, 7—8) Melecta himalayana Bingham, 1897, Fauna Brit. India, Hym. 1 : 516, fig. 172 (insect, with left wing), 2 “Kumaon, 5000 ft, and probably throughout the Himalayas at and above that altitude” Type material. — N. In dia (United Prov., Kumaon): 1 © (holotype M. himalayana Bingh.), Kumaon, North India, 4.90, Bingham coll. / Melecta himalayana Bingh. 9 Type (both labels in Bingham’s writing) (BM). Further material. — N. E. India (N. Bengal): 1 & (diss., Fig. 7-8), Kurseong, Nord-Bengal (print), Coll. Friese (print), Anthophora bengalensis Fr. [nom. nud.?}, 3, det. Friese 1913 (MNB). 1 6, Dardschiling, 14-16.X1.1929, Dr. Enslin (print), Protomelissa sauteri Friese 9 ? (in Alfken’s writing), Sig. Alfken (print). Sikkim: 1 9 (left antenna and most of abdomen missing), Sikhim, Coll. Bingham (print on yellow) (MNB). Assam: 1 & (right antenna missing), Assam, Shillong, 10. 03, Coll. Bingham (print & written on yellow), himalayana Bingh. (Bingham’s pencil writing) (ML). The holotype of this Himalayan bee was studied and redescribed by me in great detail several years before a long-anticipated journey to the Berlin museum could be realised. 274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 It was immediately recognized as a member of Cockerell’s equatorial (tropical) genus Callomelecta which, as we have seen, is congeneric with Protomelissa. Bingham’s type agrees in every respect with some additional specimens of himalayana from Northern India in the Berlin museum, listed above. The latter were probably obtained from Bingham on an exchange basis, since Alfken labelled one of them as P. sauteri, with a query. These Himalayan bees are, indeed strikingly similar to the species described by Cockerell and Friese as Melecta formosana and Protomelissa sauteri, respectively. Both are easily distinguished from other species by the bicoloured, long-fleeced thorax pubes- cence and shorn, orangish body segments. As a matter of fact, P. himalayana and for- mosana ate undoubtedly very nearly related and so closely similar to one another that they may be only geographical representatives of but one species. The pubescent colour pattern is almost the same, and the internal genital organs of the males are practically identical. Added to this, even in the shape and relative lengths of the antennal segments no differences whatsoever could be detected between them, the presence in both of a conspicuous hair-brush at the antennal scape of the male being particularly striking. Despite all this, I prefer to keep the two species apart on the basis of the characters enumerated in the key. For the rest, see the specific description of P. formosana (Ckll.). Protomelissa formosana (Cockerell) comb. nov. (Fig. 1—2, 5—6) Melecta formosana Cockerell, March 1911, Ann. Mag. Nat. Hist. (8) 7 : 227—228 (2 Kosempo, Formosa, 1908, Sauter). Anthophora sauteri Friese, 15 May 1911, Verh. Zool-bot. Ges. Wien, 61 : 127—128 (& Tainan, Formosa). — ? Strand, 1913, Suppl. Ent. 2 : 51 (9 Taihorin and Hoozan, Formosa; note). Protomelissa sauteri Friese, 1914, Dtsch. Ent. Z. : 323—324 ( 4 9 Tainan and Takao, Formosa). Type material. — Taiwan:1 @ (holotype M. formosana Ckll.), Formosa, Kos- empo, 11.08, Sauter S.V. (print on yellow), Melecta formosana Ckll. Type (in Cockerell’s hand), Type (print on light red) (MNB). 1 4, left hind leg missing (holotype A. sauteri Fr.), labelled Tainan, Formosa (written); Anthophora sauteri Fr. & det. Friese 1913 (both names in H. Friese’s writing); Type (print on dark red) (MNB). Further material. — Taiwan:1 G (diss. fig. 5—6), Formosa, Takao, 12.1908, Sauter (print), Protomelissa sauteri Fr. g', det. Friese 1913 (MNB). 2 ©, Formosa, Takao, 1908/9, Sauter (print), Protomelissa sauteri Fr. 9, det. Friese 1913 (in H. Friese’s writing) (MNB, ML). Here follows first, for completeness’ sake, the original description of Melecta for- mosana Cockerell: “©. — Length about 14 mm. Black, the abdomen stained with reddish, the hind margin of second segment broadly clear ferruginous; head and thorax with abundant long pale ochreous hair, but very long dark brown hairs on vertex and at extreme sides of face, and thorax with a broad band of dark chocolate-brown hair between the wings, also a little brown hair just below the base of the wings, and a large tuft of the same at the base of each of the very long, curved, acute scutellar spines; legs very dark reddish or reddish black, with ochreous pubescence and ferruginous spurs; hind basitarsus straight; tegulae margined with ferruginous. Wings very brown, though translucent; abdomen broad conical, without M. A. LIEFTINCK : Old World Melectine bees 275 spots, but covered with very fine appressed fox-red pubescence. Mandibles dark red except at base and apex; labrum broadly emarginate at apex; clypeus minutely granular, the linear margin shining; flagellum dark reddish; fourth antennal joint much longer than fifth, but not nearly twice as long; mesothorax shining, with strong separate punc- tures; apical plate of abdomen extremely narrow. Close to M. himalayana, Bingham, but differing in details of sculpture and pubescence. Hab. Kosempo, Formosa, 1908 (Sauter). Berlin Museum. This is an example of a Formosan species resembling a Himalayan one. The genus is new to Formosa.” General appearance of P. insidiosa (Lieft.) and vulpecula (Lieft.) (Pl. 1 Fig. 1—2), but vestiture of head and thorax still longer. Male. — Labrum (Fig. 1) light brown; disk somewhat hollowed out above, the bilobed distal portion slightly upturned, free margin thin, nearly sharp; surface rather shining, superficially reticulate-punctate, sparsely clothed with very long, raised and forward directed, silvery-yellow bristles, the fringe at free margin composed of more closely set, shorter bristles. Maxillary palpus not visible (see female). Mandibles slender, chestnut-coloured, the bases and apices dark brown; inner subapical projection small and angular, removed further distad than in P. pendleburyi (Fig. 9) and tricolor; no inner prominence near base. Antenna (Fig. 2) light to dark brown, surpassing tegula for about the combined length of two apical segments; scape robust, little curved; 2 (pedicel) retracted, annular and of minute size (not shown in Fig. 2 !); 3 in frontal view only about one-third as long as 4, the latter more than twice as long as broad (100 : 41.3); ni Fig. 1—2. Protomelissa formosana (Ckll.), & from Takao, Formosa; fig. 1, dorsal view of labrum; fig. 2, right antenna, posterior view. — Fig. 3. P. himalayana (Bingham), & from Darjeeling, dorsal view of labrum. — Fig. 4. P. tricolor spec. nov., holotype ® from Shillong, Assam, dorsal view of labrum, — Scale-lines of fig. 1, 3 and 4 (labrum), 1.0 mm, of fig. 2 (antenna), 4 mm 276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 5—13 successively shorter, from 3 on feebly longitudinally carinate, increasingly more distinctly nodose distally and clothed with minute appressed hairs, except the scape, which in front carries a dense brush of soft, golden yellow hairs. Clypeus longer and more prominent than in P. tricolor, only slightly longer than labrum, but its depth in side view more than half the greatest transverse diameter of eye (6 : 10). Eyes elongate- oval, broadest very slightly in advance of the middle, length-breadth ratio about 100 : 48; inner orbits slightly converging anterad and a little inwardly convex. Long appressed tomentum covering clypeus forming a thick pale silvery-ochreous coating, the hairs becoming long and erect posteriorly on supraclypeal area, around antennae, and on entire genal area, the hairs along eye-margin and at summit of head very long and mixed with brown. A naked, transverse band across ocellar region smooth, shining, and finely superficially punctate; occipital border again fringed with long, raised, yellow and brown hairs. Thoracic sclerites all black and closely punctate, but hidden under the dense pale pubescence, except the basal propodeal triangle, which is conspicuously naked and polished; blackish brown hair band between wing bases well defined, but its posterior limit only a short distance away from the isolated dark tufts at base of scutellar spines. Processes of scutellum long, cylindrical and tapering to a point; they are broadest at base and at first raised, but soon curve gradually backward and a little outward, the tips often rather abruptly pinched and downcurved; colour black. Legs slender, of simple structure, ferruginous to reddish brown; femora rather shining, finely superficially punc- tate, tibiae and tarsi more closely and coarsely so; outer faces of tibiae lacking spicules. Pubescence short, not very dense, except conspicuous tufts at coxae and trochanters, and long fringes at ventral ridges of fore and mid femora. Wing membrane lightly stained with ochreous, outer border of fore wing more deeply so; neuration very similar to that of P. pendleburyi (Fig. 11), but distal portion of fore wing (including marginal cell) less expanded, and second submarginal cell narrower, with its sides more closely ap- proximated costad. Integument of gaster ferruginous to reddish brown, basal portion of segments, when fully extended, dark brown, especially those of 2 and 3. All tergites rather shining basally, the punctation fine, superficial, not very close; brown appressed tomentum increasingly more conspicuous posteriorly, the hairs covering 6 and 7 hiding the surface completely. Tergite 7 tapering rapidly towards apex, which is narrow and shallowly excised. Sternites 1—5 finely punctate, the hind margins straight; 6 more projecting, strongly narrowed, the apex bluntly triangular; pubescence not dense, com- posed of appressed, decumbent, branched hairs interspersed with few long bristles, except sternite 1, which bears a tuft of long raised hairs at middle. Sternites 7 and 8 as in Fig. 5. Genital capsule 2,0 mm (incl. gonostylus); gonostylus incurved, its sides almost parallel in profile, apex not broadened, evenly rounded off; basodorsal and basoventral processes of small size, Melecta-like (Fig. 6). Female. — Resembles the male in most respects, but differs in the shape and pubes- cence of the antennae, which are equally long but more slender than in the other sex. Scape slightly curved, longer and less swollen than in male, broadest at apex (length- breadth ratio 100 : 29); long raised shock covering frontal face conspicuous, but much thinner; pedicel short and annular, though fully exposed; 3 in frontal view a little over one-third as long as 4 (but fully one-half length of 4 when measured from behind); 4 almost three times as long as broad (100 : 36), following segments successively a little shorter, but all much longer than in male and not nodose. Maxillary palpus long and slender, about equal in length to three segments of labial palpus, length of separate M. A. LIEFTINCK : Old World Melectine bees 277 segments variable (even in one individual), but first and sixth segments shortest. Body structure, colour and pubescence otherwise practically agreeing with the male, gastral sternite 5 very little prominent, its apical border almost straight. Pygidial plate black, one-third longer than its width at base (3 : 2), forming an almost isosceles triangle; margins very slightly outbent and a little thickened basally, but apex slightly produced, swollen and narrowly rounded off; surface flat almost as far as the tip, finely reticulate, apex slightly downcurved, carrying a low median ridge. Total length: $ 9 14.0—15.0 mm; fore wing 8.8—9.7 mm. The host of P. formosana is still unknown. As mentioned before, it is not unlikely that the black- and red-haired Habropoda tainanicola Strand, and/or the yellowish brown H. bucconis (Friese) 1) will prove to be the most suitable foster species of the Taiwanese Protomelissae. A nesting site of H. tainanicola was recently discovered by my friend T.C. Maa at Neihu, near Taipei, Taiwan. He came across a small colony of this during early summer of 1970, in sandy soil underneath an overhanging rock, not far away from the observer's residence. Repeated visits to this locality during April and May, 1971, yielded a second species, which, though closely similar to H. bucconis, surprisingly enough proved to be specifically distinct. The two species nested together in the same “bee village”, but the colony of H. tainanicola was the most populous of the two, albeit that the males (still undescribed !) had already disappeared. Probably this bee had established itself earlier than the new species, for individuals of either sex were present only of the latter, males even outnumbering the females. Although the nesting site was watched closely for several weeks in succession, no single Protomelissa was seen by Dr. Maa. Further observations on the spot will, it is hoped, lead to the rediscovery of the parasitic Proto- melissa formosana and its allies. Protomelissa iridescens Friese Protomelissa iridescens Friese, 1914, Dtsch. Ent. Z.: 324 (4 Takao, Formosa). — Sandhouse, 1943, Proc. U.S. Nat. Mus. 92: 592 (selected as type species of Protomelissa; not seen). The unique type of this enigmatic species could not be retraced in any collection including Friese’s main collection in the Berlin museum. The original description runs as follows: “Wie Pr. sauteri, aber Thorax ohne schwache Querbinde, Antenne erreicht nur die Flügelbasis, Abdomen mit Erzglanz. 3. Schwarz. Kopf und Thorax lang und dicht rotgelb behaart, Kopf fein runzlich punktiert, Clypeus vorgewölbt, Labrum viereckig, vorn schwach ausgerandet, Antenne rot, kürzer, erreicht nur die Flügelbasis, zweites Geisselglied viel kürzer als 3, ohne besondere Knoten. Thorax oben mit flachen, grossen Punkten, stark glänzend, aber Skulptur durch die lange Behaarung verdeckt, Scutellum jederseits mit langem, schwarzem Dorn, der aber vollkommen von der rotgelben Behaarung verdeckt ist. Abdomen fein punktiert, fast matt, schwach erzfarben schillernd, Segmente rotbraun mit schwarzer Basis und rötlich befilzt, Segmente 6—7 stark verjüngt, 7 klein und abgestutzt. Ventral- 1) Both species are represented also on the Chinese continent (Fukien) by well defined, though still undescribed, subspecies. 278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 segmente rot mit schwarzer Basis dicht punktiert, matt, mit glattem, glänzendem und häutigem Endrande, siebentes klein, gerundet. Beine schwarzbraun, gelblich behaart, Tarsen verlängert und braun, Tegulae und Flügelbasis rotgelb. L. 13 mm, Br. 415 mm. d von Takao, Sauter leg., im September 1908, Formosa (Ost-Asien).”’ Friese's statement that the “zweites Geisselglied’’ (= third segment) of the normally shaped antennae is much shorter than the fourth, would imply that the second segment (pedicel) is well visible in this species, not retracted and easily overlooked, as it is in P. formosana and himalayana. Unless the type of this puzzling bee will turn up somewhere sooner or later and prove to be congeneric with P. formosana (Ckll.), I suggest that a request be made to the International Commission on Zoological Nomenclature for action to recognize the priority of Protomelissa formosana (Ckll.) as the type of Protomelissa over Sandhouse’s unfortu- nate selection of P. iridescens Friese and give the former practical as well as de jure status by suspension of the rules. Protomelissa tricolor spec. nov. (Fig. 4) Material. — N.E. India: 1 9, Assam, Shillong, 8. 03, Coll. Bingham (print and written on yellow). Labio-maxillary complex and apical abdominal sternites dissected out and glued on card. The specimen is the holotype (MNB). Very similar to P. himalayana and formosana, but immediately distinguished from both by the form of the labrum, absence of a beard-like flocculus at the antennal scape, and by the variegated colour-pattern. Female (holotype). — Body dark brown; vertex, occiput and most of the thorax black, or almost so. Labrum (Fig. 4) short and broad, side edges obtuse-angulate, apex shallowly emarginate and a little upturned; surface slightly concave, rather shining, coarsely but not very closely punctate and clothed with strong brown bristles, which are much longer than the labrum itself and directed obliquely forward, the apical margin with dense fringe of shorter bristles. Maxillary palpus 6-segmented, slender, segment 3 longest. Mandibles strong, incurved, chestnut-coloured, but dark brown basally and growing darker also towards apex, outer border and dorsal carina with fringe of very long pale bristles; inner tooth distinct, rounded, situated slightly in advance of the apical one-fourth of mandibular length measured along dorsal carina; a second, much lower and rather crescentic prominence at about one-fifth from base. Antennae slender, not quite reaching tegulae. Scape long, slightly curved, little shorter than segments 2—4 combined, rather densely pubescent, especially in front, but hair not longer than half diameter of scape; 3 almost two times as long as its width at apex, this segment only little shorter than the following, which are subequal in length to one another and slightly less than twice as long as broad, 12 obliquely cut off and flattened beneath; 2 and 3 clothed densely with short, raised silky hairs, 4—12 practically naked, the appressed hairs being all but invisible. Clypeus about 114 times as long as labrum, surface evenly convex, surface dull, very closely punctate; entire face in front of antennae clothed with dark brown pubescence partly concealing surface and mixed with long, erect, finely branched bristle- like hairs. Eyes broader, poles more rotundate, than in himalayana and formosana M. A. LIEFTINCK: Old World Melectine bees 279 (length-breadth ratio 100 : 56), widest slightly in advance of middle in side view; inner orbits distinctly more converging anterad than in the above species and with fringe of long, dark marginal bristles. Summit of vertex, on either side behind raised frontal crest, with dense tufts of very long, erect, blackish brown feathery hairs, which are almost as long as the antennal scape. A transverse area in front and on each side of ocelli, bare, shining and sparsely punctate; vertex, occipital region as well as genal area, clothed again with dense brown pile. Thorax bulky and of great size, its entire surface including the ventral parts hidden from view by long dense pubescence composed of soft feathery hairs, which form a large subcircular, dark chocolate brown patch occupying the dorsum from the pronotum rearward to somewhat beyond level of tegulae; all around this patch the colour changes rather abruptly to canary yellow, the boundary dividing the two colours being convex posteriorly. Yellow are: the thoracic sides entirely, a narrow area around tegulae, and all of the dorsal parts behind the latter, the hairs being longest posteriorly, projecting at all sides to somewhat beyond base of abdomen, only the propodeum beneath the vestiture remaining hairless. Scutellar processes distinct, short and slightly curved, directed caudad, though barely one-fourth as long as the surrounding pubescence and completely hidden from view. Entire propodeal area polished and impunctate. Legs ferruginous, femora and bases of tibiae rather more brownish; surface smooth and rather shining. Femora finely and superficially, tibiae more coarsely and closely punctate externally, but lacking strong spicules; intero-apical process at mid tibiae long, curved and flattened; hind basitarsus subparallel-sided, very slightly outcurved. Spurs slender, slightly curved, brown. Inner ramus of fore tarsal claw half as long as outer, compressed, but little broader than this; those of mid and hind tarsi less than half length of outer, flattened and almost square. Arolia globular, of very small size. Pubescence brown, fringes at ventral ridges of femora long and rather dense, though not longer than diameter of femora, on remaining parts shorter, but more dense and silky on external faces of mid tibiae. Wing membrane yellowish, the distal border light brown, more broadly so and better defined than in himalayana; neuration as in that species, except that the second sub- marginal cell is broader, though the distal side is equally recurved. Integument of first two gastral segments dark brown above and below, tergite 2 becoming slightly more reddish brown near the end, the exposed portions of following tergites ferruginous, 3 definitely ochraceous-orange and basal areas of remaining tergites somewhat darker than their distal portions. All tergites dull, distinctly more closely punctate than in himalayana and formosana, the finely branched hairs accordingly more numerous, longest and palest in colour towards apical margins. Hind margins of sternites 1-5 shining, superficially punctate, punctures fine and sparsely distributed, much as in formosana, but pubescence shorter and less dense. Pygidial plate reddish brown, more broadly triangular than in the above-mentioned species (length and basal width in the ratio of 100 : 73.7); margins distinctly raised, sides at first a little outbent, then straight, apex narrow, almost squarely cut off; surface flat, rather shining, finely reticulate from base as far as apex, with weak indication of a short median subapical ridge. Total length: 14.0 mm approx., fore wing 9.5 mm. Male unknown. This very distinct species is the second of its genus occurring in the high mountains of northern India. 280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 5—6. Protomelissa formosana (CkIl.), 3 from Takao, Formosa; fig. 5, sternites 8 and 7, external view; fig. 6, right half of genital capsule, ventral (left) and dorsal view (vestiture partly omitted). — Fig. 7—8. P. himalayana (Bingham), & from Kurseong, N. Bengal; fig. 7, sternites 8 and 7, external view; fig. 8, right half of genital capsule, ventral (left) and dorsal view (vestiture partly omitted). All figures enlarged on the same scale M. A. LIEFTINCK : Old World Melectine bees 281 It was taken at Shillong together with P. himalayana only two months earlier than this. The two species are almost certainly parasitic on Habropoda (and maybe Elaphropoda as well). In the collection of the British Museum (Nat. Hist.) I have found a mixed series of similarly labelled H. radoszkowskii D.T., rowlandi Meade-Waldo, and turneri Ckll., all probably collected simultaneously during the summer months (August to October) of 1903, by R. E. Turner. In connection with the foregoing, it is of particular interest to note that both sexes of one of the latter, viz. H. turneri, differ from their congeners by the reduction of tarsal arolium. By a remarkable coincidence this happens to be the case also with the melectine presently described as Protomelissa tricolor, which at the same time imitates its possible host by a strikingly similar colour pattern of dark brown, light yellow and orange-red. Protomelissa habropodae (Cockerell) comb. nov. Callomelecta habropodae Cockerell, 1929, Ann. Mag. Nat. Hist. (10) 4 : 133 (4 Doi Sutep, Siam). — Lieftinck, 1944, Treubia, hors ser. : 62, 75, footnote (notes, not seen). I have not seen this species, but it is evident from the description that habropodae is congeneric with the preceding, i.e. the type species, Callomelecta pendleburyi Ckll., des- cribed a few years previously. It should be transferred accordingly to Protomelissa, with which Cockerell already had suggested an apparent relationship. The original description follows: “g. — Length about 9.5 mm. Head and thorax black, abdomen shining bright clear ferruginous, legs dusky red. Clypeus black, but densely covered with appressed pale hair; antennae brown below, practically black above, scape large and stout; flagellum not very long, barely reaching tegulae, apical joint curved; top of head, mesothorax, and scutellum polished; thorax with abundant long erect fulvous hair, not mixed with black; tegulae ferruginous. Wings brownish; marginal cell going far beyond third cubital; basal nervure going a considerable distance basad of nervulus; second cubital cell greatly contracted above, bulging on outer side, receiving recurrent nervure about the beginning of its last third. Abdomen with short appressed pale fulvous tomentum on apices of second and following segments. SIAM: Doi Sutep, on summit, Feb. 9, 1928 (Cockerell). It was flying with many Habropoda sutepensis, which it greatly resembles, and on which it is doubtless parasitic. The genus was based on a very different-looking species from the Malay Peninsula. An apparently related genus, with much longer antennae, is Protomelissa, Friese. It contains two species, occurring in Formosa.” It is worth mentioning that Habropoda sutepensis Ckll., 1929, is neither an Elaphro- poda nor a true Habropoda, as these genera are presently understood. It belongs to a group of Eurasian species nearly related to Emphoropsis Ashmead, which can probably be segregated from the latter and placed in a genus (or subgenus) of its own. This is remarkable, because the hosts of two Malaysian species of Protomelissa, as far as we know, are members of Elaphropoda. nN 00 nN TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Protomelissa pendleburyi (Cockerell) comb. nov. (Fig. 9—16) Callomelecta pendleburyi Cockerell, 1926, Ann. Mag. Nat. Hist. (9) 18 : 621—622 (gen. et spec. nov., 9 Selangor, Malaya). — Lieftinck, 1944, Treubia, hors ser. : 62—68, fig. 1—9 & 75—77, key 2 4 (incl. ö allotype, Perak and Pahang, Malaya). Hab.: Malay Peninsula. No new material. Protomelissa vulpecula (Lieftinck) comb. nov. (Pl. 1 Fig. 2) Callomelecta vulpecula Lieftinck, 1944, Treubia, hors ser. : 68—71, pl. 42 fig. 1—2 & fig. 10—16, & 75—77, key 9 4 (86 Sumatra). Hab.: Sumatra. No new material. Protomelissa insidiosa (Lieftinck) comb. nov. (PI. 1 Fig. 1) Callomelecta insidiosa Lieftinck, 1944, Treubia, hors ser. : 71—75, pl. 42 fig. 3—4, fig. 17—22, & 75—77, key 293 (96 W. Java). Further material. — Of this very rare species I collected a number of additional specimens in the mountain forests of West Java. Practically all were caught on flowers of introduced and wild Balsam (Impatiens), viz., 1 g' (topotype), W. Java, Puntjak pass, 1450 m, Telagawarna, 18.111.1953, on Impatiens chonoceras Hassk.; 9 g 1 9, W. Java, N.E. slope of Mt. Pangrango, ca. 1200 m, Tjisarua-zuid Estate, 16, 23 and 28.1V.1950, 29.V.1950, and 17.V1.1950, all in close vicinity of each other on Impatiens platypetala Lindl., in open spaces along mountain trail. A year earlier, while collecting in the same place (7.VIII.1949), a single female of Elaphropoda erratica (Lieft.) was captured, the second known example of an extremely rare species whose male still remains to be discovered. Previous collecting records at nearby Telagawarna strongly suggest a parasite-host relationship between P. insidiosa and E. erratica (see Lieftinck, 1944 : 75, and 1966 : 156). Hab.: Java. Pseudomelecta Radoszkowski, 1865 Pseudomelecta Radoszkowski, 1865, Horae Soc. Ent. Ross. 3 : 55, pl. 1 fig. 2 (insect). — Alfken, 1937, Konowia 16: 175 (key). — Linsley, 1939, Ann. Ent. Soc. Amer. 32: 434 (partim: composite key with Thyreus sp. aff. sibiricus (Rad.). — Michener, 1944, Bull. Amer. Mus. Nat. Hist. 82: 287. — Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21: 323—326 (genus restricted, diagn., distrib., figs.). — Lieftinck, 1968, Zool. Verhand. Leiden 98: 54, 56 (notes). — Osychniuk, 1970, Inst. Zool. Akad. Nauk Ukrain. SSR, 12 (4) : 49 (key). Type species: Melecta diacantha Eversmann, 1852. Prior to Popov's account (1955), this genus has been misunderstood by several authors, including Radoszkowski in his later works. Psewdomelecta in its restricted sense was redefined by Popov, who, on the basis of the original diagnosis and coloured picture, removed Pseudomelecta baerii Radosz., 1865, from it. This was thought by him to be M. A. LIEFTINCK: Old World Melectine bees 283 Fig. 9—16. Protomelissa pendleburyi (CkIl.), & and ® from Perak and Pahang, Malaya; fig. 9, right mandible of &, external view; fig. 10, hind tarsal claw of ©, caudal and lateral view; fig. 11, left fore wing of 9; fig. 12, sternite 8 of 4; fig. 13, sternite 7 of &; fig. 14, left half of genital capsule, posterodorsal view; fig. 15, right half of genital capsule, ventral view; fig. 16, pygidial plate of 9, dorsal view. After Lieftinck (1944) 284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 a true Melecta, and from an examination of the type of baerii, this indeed proves to be so. The same is true for two other melectines doubtfully referred to Psexdomelecta, viz. M. corpulenta F. Mor. and nivosa F. Mor. An analysis of these three species will, it is hoped, be given in my forthcoming revision of the genus Melecta. Popov argued that only three species should be included. These are the type species, P. diacantha (Eversm.), fuscipennis F. Mor. and atripes F. Mor. He gives good figures of the male terminalia of diacantha and new localities for fuscipennis and atripes, but of the last two no descriptions are supplied. In the following account one, P. fuscipennis transcaucasica Popov, 1955, is left out of consideration. This subspecies was based on a single female in Morawitz’ collection which came from Kazikoporan; it was poorly characterized and could not be recovered in Popov’s collection at Leningrad. As to P. atripes, 1 must refer to the original description and my own observations on the type, which are, however, too incomplete to admit of its inclusion in the key. On the present occasion two species are added to those already known, making a total of five. Never- theless, by the exclusion of atripes and transcaucasica, the absence of the male of P. chalybeia sp.n., and of the female of P. atroalba sp.n., our knowledge of the genus still remains incomplete. A peculiarity not mentioned in the description, but shared by all members, is the well pronounced limitation of the pre- and postgradular areas of the gastral tergites 2—6 (8) and 2—5 (9). The former are more strongly elevated than usual at their posterior margins, the raised graduli being convexly outbent on either side just in front of the pubescent spots so as to give the tergal surfaces a rather uneven appearence. Summarizing the key characters, the species can be distinguished from small-sized members of Melecta by a combination of the following characters: (1) partial or complete absence of pubescence on mesonotum, scutellum and para- scutella, coupled with deeply sulcate sutures between the latter and fully exposed scutellar processes; (2) closely punctate gastral tergites carrying spots which are removed far inward, away from the lateral margins of segments; (3) absence at base of gastral tergite 1 of a lateral tuft of longer, raised hairs; (4) characteristic shape of sternal plates 7 and 8 of male. Among the melectini of the Western Hemisphere, the nearctic Melectomorpha Linsley, 1939, seems to come rather near Pseudomelecta, although the few known members of the former have a different facies. By the most recent authors (Hurd, 1953), Melecto- morpha is considered a subgenus of Xeromelecta Linsley, 1939, but in the writer's opinion would better stand as a distinct genus. Pseudomelecta differs from Melecto- morpha by having unidentate (not bidentate) mandibles, longer (6-segmented) maxillary palpi, very dissimilarly formed scutellum and parascutella, and also in the more oval shape of the marginal cell, which in Melectomorpha is subtruncated at the apex. Linsley (1939) compared the type species, M. californica (Cresson), with an example of the supposed Siberian Pseudomelecta diacantha (Eversm.) which he had received from Cockerell. He arrived at the conclusion that these bees were not congeneric, in which he was certainly right. As has been pointed out by Popov (1955), however, the bee sent by Cockerell was a mis-identified species of Thyreus, probably T. sibiricus (Radosz.), or a near ally. This is evident from Linsley’s table of genera (loc. cit.: 434) in which some (not all !) of the characters given for the Asiatic Pseudomelecta are in agreement M. A. LIEFTINCK: Old World Melectine bees 285 with those of the Thyreus sibiricus group. The same species (or a near relative of T. sibiricus) was also mistaken for Pseudomelecta diacantha by Friese (1895). It must be admitted that these bees are easily confounded, resembling each other superficially in a surprising way. The following key may prove helpful to recognize the four species that could be studied in greater detail. D Sa KEY TO THE SPECIES OF Pseudomelecta No traces of white pubescent spots on abdomen, which is black, almost bare, with distinct steely blue and/or low purple reflections. Head, thorax above and legs, deep black, very shiny, clothed with erect, mainly black, pubescence not concealing surface on any part, hair longest on upper face, occipital crest, mesonotum anteriorly, and sides of metanotum. Scutellum not much swollen above, feebly biconvex, with dorsoventrally flattened, acute, triangular spines, which are slightly raised and directed straight back (Fig. 31). Legs, except femora dorsally, covered sparsely with long black hairs, lacking dense pubescent patches, surface shining. Wings clear hyaline, but distal two-thirds of fore wing membrane abruptly changing to dark brown, with slight metallic lustre. Male unknown. Hab.: Mongolia . . chalybeia . Abdomen in both sexes at least with paired white pubescent spots on gastral tergites 1—4, those at sides of 2 and 3 frequently twofold. Wings often obscured, but fore wing bases never pea changing from hyaline to dark brown. . . . 2 Males . . . Ha I TS . Females (of P. raha a CO : 5 Antenna thick, segments 4—12 transverse, distinctly broader an fone chicca distinct, transverse, crescent- or hoof-shaped. Legs black, black-haired. Upper surface of head and thorax predominantly black-haired. Distance between white lateral spots on middorsum of gastral tergites 1—4 greater than their own transverse diameter, at least so on tergites 2—4. Apex of tergite 7 narrow, subtruncated. . . . . 4 . Antenna distinctly more slender, segments 4—12, even in frontal view, squarish or a little longer than broad; rhinaria indistinct, elongate-oval, feebly impressed. Fore femur interiorly and fore tibia at base, silvery-white. Upper surface of head and thorax with much longish, silky, silvery-white and shortly branched hairs. Mesonotum, scutellum and parascutella shining, all closely punctate, but punctures much smaller than interspaces. Abdomen dull, punctation of all tergites extremely dense and fine, leaving interspaces of hardly one puncture width; all segments above clothed evenly and densely with short, appressed, finely plumose, blackish brown hairs; posterior margins bare, minutely tessellate. White spots on either side of median line on gastral tergites 1—5 very conspicuous and of large size, though not very sharply outlined, distance separating those on 1—4 on middorsum less than their own transverse diameter; spots on 1 either united to form an ill-defined transverse band, or subinterrupted medially; those on 2 and 3 twofold, the inner- most largest, oval; all dark and white hairs equally plumose. Scutellar spines strong, slightly laterally compressed and downcurved; median sulcus of concave sloping vertical area, between and behind scutellar lobes, with subtriangular white hair-spot. Fore wing membrane throughout brown with two tiny, hyaline, external spots at apical transverse veins only; hind wing subhyaline, apices slightly obscured. Tergite 7 286 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 lacking middorsal ridge, apex blunt, straight or shallowly emarginate. Structural details, Fig. 23—25. Hab.: Turkmenia; Uzbekistan. . . . . . fuscipennis Mesonotum, scutellum and parascutella shining, punctation less dense than in fusci- pennis. Scutellum small, rather pinched, strongly biconvex, with sides markedly converging, deeply hollowed out posteriorly between strongly divergent spines emerging rather abruptly from the lobes as long, slender, acuminate processes, which themselves are impunctate and shiny. Fore wing membrane brown with subhyaline streaks traversing cells lengthwise and with two confluent hyaline external spots at apical transverse veins, the free border beyond veins broadly darker brown. Ab- domen shining, all segments evenly, rather superficially punctate, punctures closer and finer than on mesonotum, but smaller than interspaces; apical tergal margins narrowly, the postgradular areas of sternites much more broadly, impunctate. Ap- pressed black pubescence covering dorsal surface of segments much sparser than in fuscipennis, the hairs not at all branched or feathery. White spots on 1—4, largest and transverse on 1, progressively smaller posteriorly, the one on 2 twofold, inner- most spot largest and transverse, outermost spot small and subcircular, as is also the spot on 4; all hairs finely plumose. Submarginal fringes at sternites 3—6 not very dense, consisting of long, suberect, blackish brown bristles. Tergite 7 lacking mid- dorsal ridge, apex blunt, shallowly emarginate. Structural details, Fig. 17—22. Hab.: Central USSR and Mongolia. . . SRE ae diacantha . Mesonotum, scutellum and parascutella jnstacless, REO a closely punctate, the interspaces barely one puncture width. Scutellum larger, main body longer with sides only slightly converging, less markedly bilobate and convex above; the lobes tapering gradually into straight, parallel, conical processes, which are more widely apart than in fuscipennis and evenly punctate. Fore wing membrane throughout very dark brown, with two minute, ill-defined clear spots at apex of outer trans- verse veins only; hind wing also darker than in diacantha. Abdomen lustreless, all segments very closely and finely reticulate-punctate, punctures much smaller, more superficial, than on mesonotum, and even more numerous than in fuscipennis. White spots on 1—4 placed much as in diacantha, the first pair subrotundate, those on 2—4 successively smaller and almost circular, the outermost spot on 2 only about one-third the size of inner and still quite some distance away from lateral border. Hind margins of sternites 2—5 shallowly emarginate medially, but more broadly so than in diacantha and fuscipennis. Tergite 7 with low but distinct middorsal ridge, apex almost squarely cut off. Structural details, = 26— 27. Female unknown. Hab.: Armenia; Transcaucasia. . . . atroalba Dorsum of abdomen dull, lustreless, Es aad very dd, dlothed with short, appressed, finely plumose, dark brown pubescence; tergites 1—4 on each side of the middle with very compact, sharply defined, snow-white subapical spots, con- sisting also of plumose hairs; spots on 1 transverse, those on 2—4 circular, but neh twofold on 2, the innermost spot then much the largest of the two. Head above, prosternum posteriorly, pronotum, mesonotum anteriorly, and scutellum on vertical surface, with rich snow-white pubescence, hairs mostly decumbent and shorter than in male; patches of white also on paraclypeal area, in front of antennae, and at base on outer face of fore tibia; sides of thorax almost bare. fuscipennis . Dorsum of abdomen shining, punctation and nature of pubescence exactly as in male: tergites covered sparsely with very short, non-plumose, black hairs; 1—4 on M. A. LIEFTINCK: Old World Melectine bees 287 each side of middle with white subapical spots consisting of plumose hairs, similar in arrangement to fuscipernis but less sharply defined, less compact; spots on 2 and 3 twofold, the outermost of these vestigial, the inner on 2—4 subrotundate, placed a little more transverse than in fwscipennis; all white hairs finely plumose. Upper part of clypeus, paraclypeal area and rest of anterior surface of head clothed with erect black hairs, those behind antennae at least as long as interocellar distance but not concealing shining punctate surface; vertex almost bare, occipital margin with long fringe of grey-brown. Thorax above mixed black and whitish, hairs more sparsely distributed and longer than in fzscipennis and predominantly white on mesonotum anteriorly; no white between scutellar lobes posteriorly. Sides of thorax also clothed with rather long suberect black hairs. Legs entirely black-haired . EE A ES OE ee diacantha The length ratios of pterostigma, marginal cell and distance separating the latter from the wing apex, are variable even in individuals of the same species. The average are for P. diacantha, 26 : 56 :100 (& ), 25 : 50 :100 (9); fuscipennis, 28 : 54 :100 (4), 20 : 47 :100 (2); atroalba, 23 : 54 : 100 ( & ); chalybeia, 27 : 50 : 100 (2). Pseudomelecta diacantha (Eversmann) (PL 1 Fig. 5; Fig. 17—22) Melecta diacantha Eversmann, 1852, Faun. Volg. Ural., Bull. Soc. Nat. Moscou, 25 : 103 (4 — not 2 ! — “terr. transuralensibus’’; “Mongolia”, teste Popov, 1955). Melecta (Pseudomelecta) diacantha, Friese, 1895, Bienen Europa's: 155 (key), 170—171 (excl. German description of 2 = Thyreus sibiricus (Rad.), teste Popov, 1955!). Pseudomelecta diacantha, Radoszkowski, 1865, Horae Soc. Ent. Ross. 3: 55—56, pl. 1 fig. 2 (4 insect) (& Transbaicalie, Irkutsk, Mongolie). — Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR, 21: 326 (discussion, distrib.), fig. 1 (wings), 21-5 (& struct.). Popov. 1960, Entom. Oboz. Moscou, 30 : 239 (note on type deposition). — Lieftinck, 1968, Zool. Verhand. Leiden, 98 : 54, 56 (notes). Type material. — Mongolia (?):1 g' (holotype M. diacantha; antennae, right fore wing, and all tibiae and tarsi of right fore and both hind legs missing), with the following pin-labels: gold disk; Radack (? print); Alp. Sibir. (print on purple); Pseudo- Melecta diacanth as, Pall. (written, Eversmann’s writing ?), ex coll. Radoszkowski Inst. Zool. P.A.N. Krakow 25/57 (print), Pseudomelecta diacantha Rad. (Radoszkowski’s writing), det. ex coll. Radoszkowski (print); Typus (written on red) (IZK). Further material. — Mongolia:1 & (right antenna and mid legs partly missing), S. Mongoliet 1927, Hutjertu Gol, Sven Hedin Exp. Ctr. Asien, Dr. Hummel (print), 218/51 (print on red), Pseudomelecta diacantha Ev. &, det. J. D. Alfken 1934 (NRS). 1 & (diss., fig. 17—18, 20—22), Mongolia, Ulan Bator, dr. Sobeslavsky, Melecta sp. d', det. Kocourek (CK). 1 9 (antennae missing), Sibir. Altai (print on red), Pseudo- melecta diacanth. (same handwriting as G holotype), ex coll. Radoszkowski Inst. Zool. P.A.N. Krakow 25/57 (print), Pseudomelecta diacantha Rad. (Radoszkowski’s writing), det. ex coll. Radoszkowski (print) (IZK). Through the kindness of Miss Dr. M. Dylewska and Dr. St. Bleszynski, I received on loan Eversmann’s type of M. diacantha, in the collection of the Zoological Institute at Krakow (see Popov, 1960). This turned out to be a male, but the same collection also contains a single example of the undescribed female, which enables me to redefine both sexes along with a few males found in other collections. 288 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Before Popov had visited Krakow to examine Eversmann’s type in Radoszkowski’s collection, he gave as the typical locality “Mongolia”, still believing the specimen to be a female. This should now be corrected, inasmuch as the type is a male, which Evers- mann had received from Pallas, originating from the “Siberian Alps’ (possibly in Mongolia) and clearly labelled “Type”. The specimen was wrongly sexed and un- questionably is the one figured by Radoszkowski, who did not even know the female at that time. According to Popov, more examples have since become known, as appears from his locality list (loc. cit., 1955 : 326), which includes more places in Mongolia and others in the U.S.S.R. (e.g. Hika, Irkutsk, Minusinsk and Tshiba, in southern Siberia). Male. — Labrum broader than long (about 4 : 3), almost flat, coarsely rugosely punctate basally, disk almost bare and polished, with few large punctures towards apex; anterior border shallowly emarginate with rounded side-angles; surface with few long black bristles, the latter more numerous and shorter at anterior border. Mandibles robust, with strong, subtriangular, blunt interior tooth at about three-fourth length from base. Tongue only little shorter than antenna. Maxillary palpus apparently 6-segmented, but last two joints fused together in dissected male from Ulan Bator (Fig. 18). Malar space smooth and shining. Antenna thick, scape curved, distinctly broadened apically, clothed sparsely with long hairs, impunctate and shining; 3 markedly longer than its width at apex but only slightly longer than 4; all flagellar segments transverse (length ratio of 3—13 as 26 : 20 : 16 : 18 : 16 : 15 :15 : 15 : 15 : 15 : 14); rhinaria on 3—13 deeply impressed, transverse, crescent-shaped, the one on 3 oval. Face moderately prominent, depth in side view more than half greatest diameter of eye (ratio 3 : 5). Clypeus and dorsal surface of head closely punctate, interspaces rather shining but barely one puncture width on clypeus and frontal area, punctures on vertex larger and more widely spaced. Pubescence black, erect and rather dense, especially on antennal area, but nowhere hiding surface, hairs about half as long as diameter of eye; vertex almost bare, but long black hairs are present on occipital area and under surface, those at occipital border partly whitish. Dorsal surface of thoracic segments shining, less closely punctate than on head, all punctures smaller than smooth interspaces, centre of mesonotum with small, almost impunctate area on either side of impressed median line at level of anterior margin of tegulae; parapsidae short, indistinct. Basal portion of tegulae coarsely striato-punctate, distally smooth. All of the thoracic sides closely rugosely punctate, punctures larger than interspaces; posterior parts behind scutellum dull, closely punctate except smooth stripes bordering propodeal triangle. Pubescence of thorax rather long and dishevelled, mostly black with whitish hairs intermixed on anterior portion of mesonotum, in front of tegulae, pure white and tufty at lateral metanotal area; at sides and underneath erect, not very dense, entirely black. Legs black, entirely black-haired; femora densely punctate, lower ridges fringed sparsely with rather long hair; outer faces of fore and hind tibiae shining, those of mid tibiae dull, the external pad dense, covering almost entire surface, hair short, very dark brown; outer face of hind tibia impressed and shining at apex, covered towards the end with short thick spicules, but margin convexly rounded and a little upturned; hind basitarsus straight and parallel-sided, dothed with long hairs; inner ramus of tarsal claws shorter than outer, on fore legs only little broader than this, but those of mid and hind legs broader, though not definitely truncated or axe-shaped, the tips being acute (Fig. 19). Marginal cell of fore wing rounded at both ends. Distal side of second submarginal cell rudimentary in left fore wing of holotype (right fore wing missing), this cell in the other males variable, occasionally very narrow, with sides almost M. A. LIEFTINCK : Old World Melectine bees 289 Fig. 17—22. Pseudomelecta diacantha (Eversm.), & from Mongolia; fig. 17, head, side view; fig. 18, maxillary palpus; fig. 19, right hind tarsal claw; fig. 20, sternites 8 and 7, external view; fig. 21, right half of genital capsule, ventral view; fig. 22, interior view of left gonostylus, more enlarged, showing ventrobasal process. Fig. 19 after 4 from Hutjertu Gol, all others after & from Ulan Bator. Fig. 20 and 21 with same scale-lines — 0.5 and 1 mm, respectively 290 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 meeting marginal cell in one point; third submarginal a little shorter than high (20 : 22). Tergite 7 gently tapering, apex rather narrow, feebly bituberculate, whole surface closely punctate and clothed with dark pubescence. Hind margin of sternites 3—5 very shallowly incurved medially; apex of sternite 6 a little produced, bluntly rounded. Sternites 7 and 8 as in Fig. 20, apex of 7 much broader than in the other species. Genital capsule (Fig. 21—22), small, ca 1.2 mm (incl. gonostylus). Female (unique). — Very similar to male, differing as follows. Labrum subquadran- gular, a little longer, length: breadth ratio of 17 : 20; anterior border straight, lacking minute subapical median tubercle. Mandibles with inner tooth robust and broad, placed more proximad, flattened with truncated apex. Maxillary palpus invisible; antennae miss- ing. Nature and colour of pubescence of head and thorax as in male, mesonotum lacking definite white spots dorsally, but hairs partly whitish at the same places as in male. Pubes- cence shorter and less dense on all parts of legs; hind tibial spurs straight and of almost equal length. Scutellar spines as in male, distance separating them about twice their length, apices slightly downcurved, not acuminate, impunctate and shining. White spots on dorsum of gastral tergites 1—4 in a regular row, those on 1 transverse. Exposed portion of gradually tapered pygidial plate convex, not carinate, dark brown, the rounded apex black. Sternites almost bare and shining; pregradular portions of sternites closely punctate, all postgradular areas broad, finely tessellate. Pseudomelecta fuscipennis (F. Mor.) (PI. 1 Fig. 34; Fig. 23—25) Melecta fuscipennis Morawitz, 1875, in Fedschenko’s Reise in Turkestan, etc, Mem. Imp. Anthrop. & Ethn. Moscou 19 : 141—142 (2 4 Zaravshan; — in deserto Kisilkum); Morawitz, 1876, Ibid. 21, tab. II figs. 26—27 (9 & insects, coloured). Pseudomelecta fuscipennis, Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR, 21: 326 (49, new locs.; excl. 2 transcaucasica subspec. nov., Zakavkazje, Kazikoporan). Type material — USSR: 1g° (diss., Fig. 23—25), lectotype M. fuscipennis, by present designation, Bairakum (print, in Russian), 4 (print on pink), Melecta fuscipennis n.sp. F. Morawitz det. 1875 (print; specific name and year in Morawitz’ writing) (ZMM). Further material. — USSR :1 3, Seravschan, Katty-Kurgan, Glasunov 1892 (print), F. Morawitz (print), fuscipennis F. Mor. g' var. (Morawitz writing) (ZIL).5 gd 8 9 (3 & wrongly sexed by Morawitz!), 2 & of these (one wrongly sexed) bearing gold disks and Sutkent (print, in Russian), fuscipennis Mor. Typ, in Morawitz’ writing, the remainder originating from Baigakum nr. Djulek, Turkestan, S. Malyshev leg., Sarafschan valley (1 9), and Katty-Kurgan, Glasunov 1892 (all ZIL). 3 &, with red labels and identified by F. Morawitz in 1875, with Sutkent Valley (in Russian), one additionally with Karak (in Russian, Morawitz’ writing) (ZMM). 1 9, 13 (print), Dzhamskoe [canyon] (in Russian), Melecta fuscipennis nov. sp. 9 (written), F. Morawitz det. 1875 (print; specific name and year in Morawitz’ writing (ZMM). 1 9, Turkestan, Sarachs, 22.6.1880, Melecta fuscipennis Mor., det. Friese 1896 (MNB). 1 9, Ferganah, Ochs, Coll. Hauser 94 (print), Pseudomelecta fuscipennis (Mor.) 9, det. D.B. Baker (NMW). The following is an almost verbatim translation of the original Russian diagnosis. “9. Black, with dark fore wings and almost smooth disk of labrum. Third antennal segment slightly longer than fourth. Head above covered with short white hair; meso- M. A. LIEFTINCK: Old World Melectine bees 291 notum anteriorly partly clothed with similar white pubescence; scutellum alongside and beneath with projecting tuft of fairly long white hair; impression between scutellar spines covered with short hair. Mesonotum and scutellum, with the exception of the afore- mentioned spots, quite bare. Prongs of scutellum acute, spine-like and rather long. Body punctation on dorsal surface fine and rather dense. Abdomen, especially the first three segments, rather densely clothed with short black tomentum by which its surface is a little shiny only along the hind margins. First four segments with two snowy-white spots situated in front of the hind borders, so that the latter as well as the side portions of the tergites remain quite black. In addition to the large spots on the second tergite, there are also two smaller spots, placed one on either side of the same segment. Legs covered with black hair. 3. Basal half of mesonotum clothed with fairly long and dense white hairs, clypeus with short silvery hairs. Second, 3rd and 4th tergites usually with 4 white spots, those on the 2nd segment frequently confluent. Segment 1 almost wholly covered with hairs; legs either entirely black, or the tibiae with white hair externally, especially on the mid tibia. Hind margins of the hindermost sternites emarginate; apical border of 7th tergite almost rounded. Somewhat resembles M. plurinotata, from which it can be distinguished by smooth disk of labrum, almost naked scutellum, lustreless (mat) abdomen, dark fore wings and shorter pubescence.” The last paragraph of the original description runs as follows: “This magnificent species is found in the valley of Zaravshan: in the Dzham canyon, 13 May; in Kara-Tube, 20 May. In the steppe Kisilkum near the ruins of Sutkent, 3 May; on Mt. Karak, 7 May, and near Bai rakum, 4 May (700—3200 ft.).” New localities mentioned by Popov were Aktasta river (Akmolinski distr.) ; Dzulek on Sir-Darja; Tshangir in Zaravshan; near Kitaba; Ashabad; and Annan. Two males, one the dissected lectotype (Pl. 1 Fig. 3), are still before me; they resemble each other closely and have the following additional characters in common. Labrum squarish, rugosely punctate at extreme base, disk almost bare and shining, apical one-third and sides more finely punctate; anterior border straight, with rounded side angles, fringed with brown bristles, the subapical tubercle narrow and distinctly raised. Mandibles with very robust interior tooth placed about midway length. Antennal scape long and strong, curved, as long as segments 3—5 united; 3 and 4 subequal in length. Pubescence surrounding pronotal tubercles and mesopleurae white, rather long but not concealing surface, hairs becoming dark brown laterally and underneath. Posterior spot of appressed white hairs between scutellar lobes; apical fringe at lower ridge and at base on outer faces of fore tibiae, white; for the rest legs are dark brown lacking white hairs. Outer faces of mid tibiae at extreme tip, as well as most of the external faces of the hinder pair, smooth and shining, the latter covered with numerous sharp spicules. Inner rami of all tarsal claws broad, flattened and axe-shaped. Marginal cell of fore wing almost to fully three times as long as broad, apex rounded. Third submarginal cell almost as long as high; sides of second submarginal strongly converging anteriorly, fused to- gether and shortly stalked before reaching anal side of marginal cell in both fore wings of the lectotype. White postgradular markings on gastral tergites 1—5 approximated middorsally, as described in the key, those on 4 and 5 more widely separated, on 5 very small and often wanting; spots on 1 frequently united and band-like, but subinterrupted 292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 23—25. Pseudomelecta fuscipennis (Mor.), & lectotype from Bairakum; fig. 23, tergite 7, ventral view; fig. 24, sternites 8 and 7, external view; fig. 25, right half of genital capsule, ventral (left) and dorsal view. Fig. 24 same magnification as fig. 26 of P. atroalba posteriorly in the lectotype (Pl. 1 Fig. 3). Hind margins of tergites 4—6 more broadly impunctate than preceding ones, their surface finely tessellate and rather shining; apex of tergite 7 blunt, almost squarely cut off or shallowly emarginate, whole surface clothed with dark hairs (Fig. 23). Exposed basal areas of sternites 2—6 closely punctate, less densely brushy than in P. diacantha, the dark hairs not exceeding hind margins of sternites; distal areas rather shining, finely tessellate; hind margin of sternite 2 very slightly and narrowly, of 3—5 much more broadly and deeply emarginate, distinctly crescent-shaped. Apex of sternite 6 scarcely produced, its margin almost straight. Sternites 7 and 8 as in Fig. 24. Genital capsule (fig. 25), ca 1.5 mm long (incl. gonostylus). Female (Pl. 1 Fig. 4). — Averages smaller in size than male. Labrum subquadrangular, slightly shorter than its greatest width across middle (17 : 20); basal portion slightly convex, with few punctures, then flat, rugosely punctate, with marginal fringe of strong black bristles; anterior border shallowly emarginate, the subapical median tubercle of minute size. Face little projecting, depth in side view less than half diameter of eye. Clypeus convex, surface slightly shining, closely finely punctate, interspaces about one puncture width, clothed with decumbent dark brown pubescence at base, for the rest hairless. Punctation of dorsal thoracic segments and abdomen much as in male, but punctures even more numerous on all parts, the dorsal surface and sides of thorax accor- dingly less shining. Integument of legs and abdomen dark reddish brown, not quite black as in diacantha; pubescence of legs shorter and less dense than in that species. Shape of submarginal cells variable, but second cell invariably broader than in P. diacantha, the fractured anal side longer (as in male); third submarginal distinctly shorter than high (15 : 21). Abdomen compact, entirely lustreless above, the white pubescent spots much more sharply outlined than in P. diacantha and often almost circular; postgradular areas M. A. Lrertinck: Old World Melectine bees 293 of sternites 1—4 broadly impunctate, all of the exposed surface of 5 and 6 closely punctate. Pygidial plate gradually tapered, with gently rounded apex; surface (except at extreme base) convex, with low median ridge, finely tessellate, side-margins subacute and a little upturned almost as far as apex; colour dark reddish brown, bordered with black. Measurements variable. Total length: $ 12.0—12.5 mm, fore wing 9.0 mm; 9 9.0—10.5 mm and 7.5—9.0 mm, respectively. Hab.: Southern USSR: Turkmenia; Uzbekistan. In January, 1965, Dr. A. N. Zhelokhovtsev kindly sent to me for inspection the supposed types of this species, a male from Bairakum, here selected lectotype, and a female from Dzhamskoe Canyon, which may be considered lectoallotype, both labelled M. fuscipennis by F. Morawitz himself and deposited in the collection of the Zoological Museum at Moscow. These specimens were compared by me in 1968 at Moscow with red-labelled males from Sutkent standing over a drawer label M. corpulenta, but also carrying identification labels M. fuscipennis, written by Morawitz in 1875. These pinned individuals, along with some others in the Leningrad museum, are undoubtedly con- specific. Although a number of the latter bear witness of having been studied by V. B. Popov, it is unfortunate that none of them were determined by him, and I have also looked in vain for the female from Kazikoporan, on which Popov based the new sub- specific name transcaucasica. Morawitz’ coloured illustrations of the two sexes of this bee give a good impression of the colour design, although the general aspect of both male and female is more dumpy than would appear from these pictures. Pseudomelecta atroalba spec. nov. (Fig. 26—27) Material. — USSR: 1 & (diss, Fig. 26—27, holotype), Transcaucasia, labelled Eriwan, 1898, Korb (print), 17 (written) (MNB); 1 3 (paratype), with same locality label and Melecta fuscipennis Mor. Kauk., in H. Friese’s writing (MNB); 1 3 (para- type), Asia min. 1890 (print), 17 (written), Melecta fuscipennis Mor. d', det. H. Friese 1900 (Friese’s writing) (ML). Male. — Labrum black, little broader than long (27 : 22), basal tubercles convex, smooth, with few deep punctures; median area of disk impunctate, irregularly rugose (in one & with few coarse punctures), this area somewhat triangularly pointed, ending in a low marginal tubercle; rest of surface closely punctate. Disk of labrum with few raised hairs that become longer and more numerous towards anterior border, which is almost straight. Mandibles with strong, incurved and pointed inner tooth slightly beyond halfway length; bases with numerous closely set fine punctures. Maxillary palpus 6- segmented, 2 and 6 longest, both distinctly longer than each of the intermediate segments. Malar space linear, shining. Antenna shaped similarly to P. diacantha, but still thicker, the curved scape shorter and more broadened apically, ratio of length and width at apex 100 : 48 (diacantha, 100 : 40; fuscipennis, 100 : 31.6); segment 3 only a trifle longer than succeeding segments; rhinaria as in diacantha. Face much as in P. diacantha, but clypeus more convex. Whole anterior and dorsal surface of head closely deeply punctate, punctures on clypeus, paraclypeal and frontal areas separated by less than one 294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 26—27. Pseudomelecta atroalba spec. nov., & holotype from Erewan; fig. 26, sternites 8 and 7, external view; fig. 27, right half of genital capsule, ventral (left) and dorsal view (ventrobasal process shown in both aspects: dorsobasal process wanting). Scale-line = 0.5 mm puncture width on dull ground; surface of vertex more shining, with smooth impunctate and impressed areas on each side of lateral ocelli. Punctures more closely set and larger than in both diacantha and fuscipennis. Pubescence on upper face and antennal area deep black, the erect tufts behind antennae long and dense, hiding the surface from view, behind which the pile becomes sparse, decumbent and mainly silvery, notably on vertex, postoccipital area and occiput; genal area clothed with long, raised black hairs. Texture of thorax as described in the key; all of the sides and posterior parts behind scutellum closely, rugosely punctate, lacking smooth sutural stripes. Pubescence short, sparse and only partly raised, mainly black; thin patches and narrow lines of white decumbent hairs forming indistinct more or less confluent ms-/pn-als-plsa spots, are present on dorsal surface; lateral metanotal area and posterior surface between scutellar lobes also shortly white-haired in the holotype (black in the others). Legs black, entirely black- haired; texture and pubescence quite similar to diacantha, except that the inner rami of all tarsal claws are in the form of very broad and thin plates, larger and more distinctly axe-shaped than in that species, and even larger than in fwscipennis. Venation of fore wing similar to fzscipennis, sides of second submarginal cell strongly converging anterad, meeting marginal cell in a single point, or almost so; third submarginal about as long as high. Abdomen short, thick-set, broadly oval, completely lustreless above. Pure white tergal markings on 1—4 only, all spots sharply defined, in a regular row, those on 2 twofold, the outermost of these very small; dark ground on either side of (innermost) spots on 2—4 well visible in dorsal view, as in the allied species. Sternites all black, black-haired; surface densely punctate, the apical strips rather broad, smooth and finely tessellate, the margins very shallowly emarginate at middle; bristles shorter than in M. A. LIEFTINCK: Old World Melectine bees 295 diacantha. Apex of sternite 6 an almost rectangular triangle. Sternites 7 and 8 as in Fig. 26. Genital capsule (Fig. 27) small, ca 1.3 mm (incl. gonostylus). Total length: 11.5—12.5 mm, fore wing 9.2—9.5 mm. Female unknown. A stockily built species, with a large, globular thorax and very dark wings. Distinctive features are the compact ovoid shape of the abdomen, coarsely punctate, dullish black integument, and almost complete absence of white pubescent markings on head and thorax. The sharply defined white abdominal spots are placed much as in the female of fuscipennis, but the male of the latter is more profusely white-haired, has much slenderer antennae, and lighter wings. In some respects the new species comes nearer diacantha, although this is a more shiny insect, in which the antennae and legs are less swollen than in atroalba. P. atroalba is the most westerly distributed member of the genus. Pseudomelecta atripes (F. Mor.) Melecta atripes Morawitz, 1895, Horae Soc. Ent. Ross. 29 : 40—41 (4 Tschuli, Turcmenia). Pseudomelecta atripes, Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21: 326—327 (& Topjatan, Uzboi). Material. —USSR:1 4, Perevan (transl. form Russian), 26.1V.1889, with illegible writing, coll. Morawitz (print), Melecta atripes F. Mor. & (in Morawitz’ hand) (ZIL). 1 g, illegible locality (probably Topjatan, Uzboi, leg. D.M. Steinberg, 24.1V.1951, on Astragalus, as given by Popov, 1955), Pseudomelecta atripes F. Mor., det. V. B. Popov (ZIL). Original description. — “ 9 mm. Bei diesem Männchen sind Kopf und Clypeus, das Gesicht, das vordere Drittel des Mesonotum und der obere Teil der Mesopleuren weiss- lich behaart; diese sind fein und dicht runzlig, jenes nebst dem fast kahlen Schildchen spärlicher punktirt, indem die glänzenden Zwischenräume der Punkte meist breiter als diese sind. Zu beiden Seiten des Schildchens steht ein Busch weisser Haare. Die Seiten des Mittelsegmentes, die Brust, die Bauchringe und die Beine sind einfarbig kohlschwarz behaart. Die erste Dorsalplatte des Abdomens hat jederseits eine querovale weisse Haar- makel; auf der 2-ten, 3-ten und 4-ten ist zu beiden Seiten der Scheibe, dem Endrande näher stehend, eine rundliche weisse Haarmakel vorhanden; ausserdem sind noch auf der 2-ten und 3-ten nahe dem Seitenrande sehr kleine, aus wenigen weissen Haaren zu- sammengesetzte Flecken zu sehen. Die glänzenden Flügelschuppen sind schwarz. Die pechschwarz geäderten Flügel sind dunkelbraun getrübt. M. fuscipennis F. Mor. ähnlich; diese ist aber grösser, deren erstes Abdominalsegment fast vollständig weiss befilzt; die weissen Haarmakeln der folgenden sind, namentlich die am Seitenrande stehenden, grösser, fliessen auf dem zweiten zusammen; die Schienen des mittleren Beinpaares sind aussen vollständig, die des vorderen am Grunde weiss befilzt. Bei Tschuli von A. Semenow gesammelt.” Popov (1955 : 326) rightly says that the specimens in the Leningrad Museum do not fully agree with the description, the localities being also different. Unfortunately, my observations on this species, made while studying bees in the col- lections of the Leningrad museum in 1968, are incomplete. However, on comparing the 296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 above males with those of P. diacantha and fuscipennis, I noted the following. A true Psendomelecta. Very similar superficially to certain Central Asiatic Thyreus, but apart from the primary generic characters separating them (e.g., the relative lengths of the first two gastral segments), distinguished by decumbent and at the same time denser and longer white pubescence upon the anterior portion of mesonotum, the deeply concave area between the scutellar spines, and white gastral spots consisting of appressed tomentum. Scutellar processes rather long, directed straight back. Similar to P. diacantha, but body of the latter more compact, scutellar spines more closely approximated, the area separating them deeper. Also similar to P. fuscipennis, which likewise has a more thick-set body than atripes, shorter pubescence and more widely distant scutellar processes, the area between the latter being more densely clothed with felt-like hairs. The antennae and genital organs of the holotype male will have to be studied before anything definite can be said about the position of this species. In any case it is a much slenderer, more Thryreus-like insect than our new species P. atroalba, described in the previous pages. Hab.: Turkmenia. Pseudomelecta chalybeia spec. nov. (PI. 1 Fig. 6; Fig. 28—32) Material. — Mongolia: 9 9, S.W. Mongolia, Aimuk Bajanchongor: N. of Altai-Gobi, S.E. shore of lake Bun-cagan-nur, 7.VI.1962 (1 ex.); Transaltai-Gobi, S. of Argalant Mts., Somon Bajancagan, 9.VI.1962 (6 ex.); same area, oasis Ouro Dzun-mod, 12.V1.1962 (1 ex.); all R. Piechocki, Mongolisch-Deutsche Biologische Expedition 1962. Holotype, Somon Bajancagan, 9.VI.1962, and four paratypes from the above localities (MHW); three paratypes (ML), one paratype (USNM). Female. — General appearance (Pl. 1 Fig. 6). Labrum deep black, squarish (Fig. 29), rugosely punctate all along extreme base, basal tubercles poorly defined, centred with brown; disk flat, median portion subtriangular in outline, ending in a minute marginal tubercle, surface bare, polished and very shining, with a few scattered punctures; side portions irregularly, closely punctate, sparsely clothed with raised black bristles and a dense comb of shorter hairs fringing anterior margin, the latter shallowly and broadly excavated with a few minute crenations. Mandibles with strong, blunt inner tooth placed at three-fifths to three-fourths distance from base; bases with numerous minute, super- ficial punctures externally. Malar space linear, surface polished. Proboscis (labio-maxillary complex) when fully extended, longer than antenna (ratio about 100 : 71). Maxillary palpus 6-segmented, 1 very short, 2—4 slender, little more than five times as long as broad and subequal in length, 5 about two-thirds and 6 only half as long as preceding segments. Antenna shaped much as in diacantha female, but scape slenderer, a little more than three times as long as its width at apex (100 : 30) and about as long as 3—5 united, clothed sparsely with long black bristles above and below; segments 3—12 all a little longer than broad and subequal in length to one another. Eyes elongate-oval (Fig. 28). Face projecting, depth about two-thirds greatest diameter of eye (Fig. 28); clypeus perfectly flat above, side-edges obtuse-angulate, whole surface shining, striato- punctate, punctures of different sizes, much smaller than interspaces anteriorly, but becoming more numerous upward; frons, vertex, occiput and genal area closely punctate, M. A. LIEFTINCK : Old World Melectine bees 297 the interspaces hardly one puncture width; a crescent-shaped polished area on either side of lateral ocelli. Pubescence of head scanty, rather long, erect and black on upper facial parts, becoming still longer and denser between and behind antennae, the tufts behind antennae consisting of silky, pure white and finely branched hairs; ocellar area almost bare, but occipital area again fringed with longish white hairs; genal area black. Mesonotum with distinctly impressed median and parapsidal lines, the latter not ex- ceeding tegulae in length; surface smooth, very shining, punctation rather deep and irregular, with polished, almost impunctate middorsal areas running lengthwise on either side of median line and also lateral to parapsidal lines, the punctation becoming very dense, leaving no interspaces, anteriorly and at sides of mesonotum. Bases of tegulae with punctures of different sizes. Parascutella more coarsely and closely punctate than scutellum, the surface of which is rugose, partly reticulate-punctate; scutellar spines flattened dorsoventrally, upturned and hollowed out above, with almost impunctate and shining apices (Fig. 31). Metanotum finely rugose, lustreless, the propodeal triangle and lateral parts of same rugosely punctate with slightly shining sutural lines; sides and under surface of thorax throughout closely, very coarsely punctate, the punctures larger and deeper than those on postscutellar parts. Black thoracic pubescence scanty, hairs partly raised, resembling pile of diacantha, dorsal surface almost naked, but semierect white hairs longer, forming a pro-mesonotal collar that extends back to as far as base of tegulae; also small tufts of white around pronotal tubercles, at base of parascutella, and longer tufts of white on lateral metanotal area; raised hairs behind scutellum and at sides of thorax shorter, not hiding surface, black. Legs slender, normal; coxae and femora shining, sparsely superficially punctate; armature and pubescence of tibiae and tarsi as in diacantha; inner rami of all tarsal claws distinctly broadened, very similar to the last-mentioned species, but shorter and less definitely axe-shaped than in atroalba (Fig. 30). Wing venation variable; sides of second submarginal cell of fore wing less strongly converging towards marginal cell than in the other species, the proximal side often straight and distal side in three out of the total incompletely developed; in most females the first recurrent vein approaches the distal side of the second submarginal more closely than in the allied species, and in both wings of one female these veins are interstitial. Third submarginal also variable in shape, but always a little higher than long and occasionally subequal in size to the second. Abdomen shining, all tergites bare, with distinct, though low, metallic blue lustre, the pregradular areas occasionally more greenish and postgradular areas with purplish hue; graduli of 2—5 well indicated, slightly raised, especially at the sides, those of 2—4 distinctly convex posteriorly; all segments closely, finely and superficially punctate, with series of larger punctures intermixed at the graduli; all punctures smaller than interspaces; surface of postgradular areas of 4 and 5 finely tessellate and almost impunctate towards apical margins. Pygidial plate rather broad, surface flat, finely tessellate, apex with distinct, broad and smooth, median ridge (Fig. 32). Sternal surfaces shining and with few longish bristles fringing the undulated boundary between gradular areas; pregradular portions closely punctate on low metallic blue or green ground and with irregular row of large punctures separating them from the distal portions, which are broad, smooth, impunctate and widest at middle, with slight coppery reflections; hind margins of sternite 1 straight, those of 2—5 very slightly concave; sternite 6 finely striatopunctate. Total length: 10.5—11.0 mm, fore wing 8.5—9.0 mm. Male unknown. 298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 28—32. Pseudomelecta chalybeia spec. nov., ® paratype from Mongolia; fig. 28, right side view of head; fig. 29, dorsal view of labrum; fig. 30, mid tarsal claw, right external and caudal view, setae omitted; fig. 31, scutellar area, right lateral and dorsal view; fig. 32, pygidial plate, dorsal view This extraordinary species, by its metallic-coloured body and sharply defined wing markings, takes an isolated position among all other melectines of the Eastern Hemi- sphere. It has no near allies, not even among its congeners, and although it is here placed in Pseudomelecta with some misgivings, I can see no sufficient grounds yet for establishing a separate genus to hold it. The absence of a male is unfortunate, for this may reveal characters by which the species differs more appreciably from the other members of the genus. Nothing, of course, can be said about its host, but it may be mentioned in passing that M. A. LIEFTINCK : Old World Melectine bees 299 a metallic coloured Anthophora potanini has been described also from Mongolia by Morawitz (1890, Horae Soc. Ent. Ross. 24 : 353) after a single female measuring 12—13 mm. A second metallic species of the same genus, viz. A. aeneiventris Hedicke, is known from the North Pamir Mts. This is likewise of small size, the female described measuring 10.5—11 mm (Hedicke, 1931, Mitt. Zool. Mus. Berlin, 16 : 850—851). Paracrocisa Alfken, 1937 Paracrocisa Alfken, 1937, Konowia 16 : 172—175. — Sandhouse, 1943, Proc. U.S. Nat. Mus. 92 : 583. — Popov, 1955, Trudi Zool. Inst. Akad. Nauk USSR 21 : 322, 327—329, redefined and amplified, with fig. 2 (fore wing) and 3 (& terminalia), of P. glasunowi (F. Mor.). — Osychniuk, 1970, Inst. Zool. Akad. Nauk Ukrain. SSR, 12 (4): 49 (key), fig. 23° (fore wing). Type species: P. sinaitica Alfken, 1937, ® & S.W. Sinai Peninsula. Paracrocisa was proposed for the reception of a single pair of a peculiar melectine, the proper status of which has long remained a mystery. Owing to the shockingly inadequate generic diagnosis, and of the description of its type species as well, no sub- sequent records of P. sinaitica are to be found in the literature. The identity of the female “Typus” is still uncertain; it was stated by Alfken to be in Kustos A. Kneucker's collection and may have been lost, but the male “‘Allotypus’’, selected as such by Alfken, fortunately remained in his private collection. After Alfken’s death, this bee came into the possession of the Berlin Museum (MNB). In 1970, while going over Alfken’s bees at Berlin with Dr. Königsmann, I finally recovered the specimen in question, whereupon allowance was made to study it in greater detail. This male is here selected lectotype, bearing in mind that a comparison with the type of the much earlier described P. guilochei has revealed that the two are very nearly related and may even prove to be conspecific. Popov (1955) lists five species, but P. caesareae (Friese), which he knew only from the description, was wrongly assigned to Paracrocisa; it is, in fact, an aberrant member of Melecta, whose members will be discussed in a later part of this revision. I have accepted Popov’s re-characterization of the genus, as this is based on the nearly related P. glasunowi (F. Mor.), at that time the only species with which he was actually acquainted. As we will see, P. glasunowi is synonymous with the earlier defined P. kuschakewiczi (Ra- doszk.). This leaves us with three closely allied species, which will be discussed hereafter. Apart from the generic characters mentioned in the key, the following diagnosis may serve to recognize fresh examples of either sex. Head. — Labrum about three-fourths as long as its greatest breadth at middle; basal tubercles low but of large size; surface hollowed out medially, coarsely rugosely punctate, its distal half clothed sparsely with raised, usually white, hair; anterior border straight, but margin a little thickened, with small, upturned median tubercle (Fig. 33). Mandible- bases of male often with white hair-spot. Clypeus short and broad, surface convex, very finely reticulate-punctate and in male clothed densely with decumbent, snowy-white, silky pubescence, the hairs becoming longer and upright posteriorly to as far as the level of ocelli; this pubescence much shorter and finer in female, especially on clypeus and supra- clypeal area. Vertex closely punctate, except slightly impressed, shiny hairless areas on either side of lateral ocelli. Whole occipital and upper portion of postocular area with raised white hairs, longest in male. Antennal scape with very short appressed white hair in front; flagellum dull anteriorly, smooth and more shining posteriorly. 300 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Thorax. — Dorsum and sides strongly, closely punctate, punctures rather deep and often coalescent, especially on scutellum. This convex, strongly declivous between the processes; the latter varying in length, at most one-third length of main body, parallel or slightly divergent, usually spine-like and pointed. Parascutella triangular, likewise convex. Surface of all posterior thoracic segments including propodeum dull, rather coarsely striato-punctate, lacking smooth areas. Tegulae finely punctate. Male with com- plete, broad, mesothoracic collar composed of semierect white hairs occupying all of the sides, except almost smooth metepisternum and a small area above mesocoxae; appressed patches on pls, longish tufts behind wings and on /p. Female lacking mesothoracic collar of longish hair, but instead with isolated spots /px-als, ms, pls, confluent hypm- deps, and long posterior tufts as in male, the lower portion of the thoracic sides and venter remaining black-haired. Legs. — Femora laterally compressed in both sexes, posterior longitudinal ridges obtuse; surface shining, finely superficially punctate, almost naked; outer faces of mid and hind tibiae coarsely rugose, with smooth shining interspaces and beset with strong acute spicules; outer face of hind tibia moreover with small, impressed subapical area. Wings. — Submarginal cells of fore wing shaped similarly in all species, the area occupied by the first and third being almost the same, the third only little longer than high. Also, the length ratio of pterostigma, marginal cell and distance separating the latter from the apex, differs little among the species (e.g., kaschakewiczi, 2.4 : 6.7 : 10; sinaitica, 2.2 : 6.1 : 10; guilochei, 3.0 : 7.6 : 10). Abdomen. — Gastral tergites 1—5 or 1—6 (male) and 1—4 (female) with lateral patches of appressed white pubescence, those of 1 extending from base to apex; sternites predominantly black-haired. Tergite 7 of male blunt, apex squarely cut off or shallowly emarginate with slightly raised, rounded side-angles; surface finely reticulate-punctate, clothed with dark appressed pubescence. Tips of dense brushy hairs fringing sternites 3—5 of male slightly upcurved. The apical sternal plates and genital organs of the male were first described by Radoszkowski, who also noted the peculiar appendage described below as the ,,mem- branous flap”, at the base of the gonostylus. This structure was considered by Radosz- kowski to represent the volsella. He writes: “...... la véritable volsella du forceps n'est pas soudée avec sa branche comme chez la Crocisa et la Melecta, mais est detachée et forme une partie indépendante”, the last observation being quite correct. Popov also gave sketches of the genitalia, but in his drawing (loc. cit., fig. 35), the “volsella” is shown to form part of the gonostylus, which is not the case. I am unable to corroborate Radoszkowski’s interpretation of the volsella. The appendages of the genital capsule are probably manifold and may be exemplified for P. kuschakewiczi, as follows. Capsule of large size (2.8 mm, incl. gonostylus); general appearance in ventral and dorsal aspect as in Fig. 41, 43, 45. In ventral view, the gonocoxite carries at its extreme base a pair of small, roundish interobasal lobes, which are closely approximated and placed nearest the median line, forming sclerotised, naked tubercles. Along its entire hind margin, the gonocoxite is bordered by an elongate, semitransparent thin membrane, which extends outward until meeting (and partly overlapping) a heavily sclerotised tubercle, placed on the inside at the base of each gonostylus, this tubercle bearing a tuft of thick, strong bristles. In dorsal view, the gonocoxite is seen to bear three appendages, the smallest of these being an interobasal gonocoxal tubercle carrying a dense apical hair tuft, followed beyond the capsule by a very conspicuous semitransparent scale, which M. A. LIEFTINCK: Old World Melectine bees 301 arises from the inside of the apical margin of the gonocoxite; at its base this parchment- like, incurved plate is partly covered by the broad stem of the gonostylus, but remains completely free from it; the whole surface of this membranous flap is coated with rather long plumose hairs, a dense row of similar fine setulae fringing also its entire margin. In lateral view, the gonostylus is very broad basally, for the greater part membranous, semitransparent and strongly convex externally, forming a naked subtriangular shield, of which only the outer border and narrow apex are darkly pigmented and sclerotised; in dorsal view, the styli are slenderly lanceolate, twisted and incurved, their basal portions almost bare and the rounded tips tufted with long bristles (Fig. 41, 43, 45). The hosts of Paracrocisa are unknown and we can only guess at their identity. Popov (1955) mentions Anthophora (Paramegilla) semperi Mor. as the possible foster species. This is not unlikely, because the distribution and phenology of the latter harmonize with those of P. glasunowi (= kuschakewiczi Radoszk.) in Turkestan and Kazakhstan. The supposed host is a conspicuous white-spotted bee of about the same size, superficially resembling a Melecta in certain respects. KEY TO THE SPECIES OF Paracrocisa 1. Male and female. Integument dark reddish brown, more rarely deep black. Parapsidal and median mesonotal lines markedly furrowed, especially anteriorly. Abdomen almost lustreless, all segments finely tessellate, very closely and super- ficially punctate, the interspaces about one puncture width; dark pubescence dense, consisting of minute, appressed, finely branched hairs. Male. Apical process at lower margin of hind tibia short, oval and a little upturned, apex hollowed out within, armed with marginal comb of 4—7 short, strong spicules; S-shaped inner hind tibial spur fully twice as long as gently curved outer spur. Lower margin of hind basitarsus without small subbasal prominence, but clothed instead with small patch of appressed, shining, brown hairs (Fig. 34). Subapical brush on gastral sternites 3—5 moderately long, tips of bristle-like hairs not projecting beyond hind margin of sternites. Sternite 6 abruptly narrowed, its apex a little protuberant but hind margin almost truncated; surface slightly impressed, densely pubescent, post- gradular area clothed with longer hairs. Apex of sternite 8 bilobed and distinctly emarginate (Fig. 40). Pubescent flap at base of gonostylus more or less trapezoidal, hind margin broadly and shallowly emarginate (Fig. 41). Pygidial plate of female narrowly triangular, almost two and a half times as long as its width at base; sides at first converging and slightly incurved, almost parallel, margins raised and carinated; disk flat, finely tessellate, reddish brown, tip obscured, distinctly swollen, its surface smooth. Hab.: from Turkey to E. Kazakhstan . . . kuschakewiczi — . Male and female. Integument deep black. Parapsidal and median mesonotal lines only slightly impressed. Abdomen slightly more shining, punctation fine but punctures less numerous, separated by more than one puncture width; dark pubes- cence accordingly less dense, but otherwise similar to preceding species. Male. Apical process at lower margin of hind tibia longer, more slender and slightly twisted, almost three-fourths length of outer spur, apex carrying 3—5 short, strong spicules; S-shaped inner hind tibial spur less than twice as long as gently curved outer spur. Lower margin of hind basitarsus with small, roundish subbasal prominence, the surface of which is polished and naked (Fig. 36). Subapical brush on sternites 3—5 302 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 more conspicuous, composed of thick bristles, the apical ones comb-like with tips projecting markedly beyond hind margin of sternites. Sternite 6 more evenly and broadly rounded, apex only slightly projecting, hairs longer, thicker and more definitely plumose than in kuschakewiczi. Apex of sternite 8 squarely cut off, at least slightly produced medially (Fig. 42, 44). Pubescent flap at base of gonostylus narrower, deeply trilobate (Fig. 43, 45). Pygidial plate of female as in kuschakewiczi, but tapering more gradually, sides straight with raised margins; apex scarcely swollen, with distinct low median ridge. Hab.: from Israel westward to Morocco guilochei and sinaitica Paracrocisa kuschakewiczi (Radoszk.) (Pl. 2 Fig. 10; Fig. 33—35, 40—41) Pseudomelecta kuschakewiczi Radoszkowski, 1890, Hor. Soc. Ent. Ross. 25 : 247—248, pl. II fig. 2—3 (& Vernoé — ? Alma Ata sec. Popov, 1955). Radoszkowski, 1893, Bull. Soc. Imp. Nat. Moscou, n.s. 7 (2—3) : 185, pl. VII fig. 41a-c, i & k, & sternal plates & genit. (misspelled kuchakevitzi). Paracrocisa kuschakewiczi, Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21: 329 (not seen; transferred to Paracrocisa, and compar. notes with P. glasunowi (F. Mor.). Popov, 1960, Entom. Oboz. Akad. Nauk USSR, 39 : 237—240. Melecta glasunowi Morawitz, 1895, Hor. Soc. Ent. Ross. 29 : 41—43 (9 Koschlagar and Pul-i-Chatun, S. Turcmenia). — Syn. nov. Paracrocisa glasunovi (sic), Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21: fig. 2, fore wing and 31-7, & struct. 9 &, transferred to Paracrocisa; compar. notes with P. kuschakewiczi (Rad.); addit. descr., distrib., USSR and Iran. Type material. — USSR: 1 ©, Uzbekistan, Katta-kurgan, 20.VI.1931, V. Gussa- kovski (written, transl. from Russian), P. glasunovi Mor., det. V. B. Popov; neotype Ps. kuschakewiczi Rad., by present designation (ZIL). 1 9, Southern Turcmenia, labelled Pul-i-Chatun, Melecta Glasunowi F. Mor. ©, in F. Morawitz’ writing; Coll. Morawitz (print) ; lectotype Melecta glasunowi F. Mor. by present selection (ZIL). Further material — USSR: ig 19, Transcaspia, F. Morawitz (in Alfken’s writing), both with red label Paratypus (print) and Melecta Glasunovi Mor. ab auct. 1904 (in Alfken's writing), d with additional label Pseudomelecta kuschakewiczi Rad., Mor. sp. ist synonym ? (in Alfken’s writing), Slg. Alfken (print) (MNB); 1 9, Turkestan, Pul-i-Chatun/glasunovi F. Mor. (in Morawitz’ hand) (MNB); 1 2, Turc- menia, Gaudan Pass, 5th feet, Transcaspian region, Filipovich 1897 (print, transl. from Russian), det. V. B. Popov (ZIL); 1 & (diss, PI. 2 Fig. 10 & Fig. 40—41), Turcmenia, ex coll. Morawitz, Melecta sp.? Pseudomelecta, det. Friese 1893, Paracrocisa sp. aff. glasunovi Mor., det. D. B. Baker (NMW); 1 9, Russ. merid., Elisabethpol, Kinder- mann, P. glasunowi Mor., det. D. B. Baker (NMW). Iran: 10, N. Persia, Shakhrud, 26.V.1914, Kirichenko (print, transl. from Russian), P. glasunovi Mor., det. V. B. Popov (ZIL). Afghanistan: 1 & (diss, fig. 33—35), N. Afghanistan, Prov. Herat, Wadi-i-Namak Sor, 400 m, 16.VI.1964, coll. O. Jakes (BRNO). Asia minor (Tur- key): 1 g 1 9, Asia min. 1890, both with Melecta baerii Rad., det. Friese 1900, the 3 with additional labels Melecta niveipes ! & 2 det. Enderlein, and Melecta baerii Rad. (Friese’s hand) (MNB, ML); 1 9, O.-Anatolien, Eski Malatia, 5.7.1937, Ramme leg. (MNB); 1 9, Asia min., Naday 1911, Dinek, VI.1926 (MBUD); 1 9, Turkei, Konya, 23.V11.1971, Kl. Warncke (CW). M. A. LIEFTINCK: Old World Melectine bees 303 As mentioned above, the copulatory organs of the unique type of this species were dissected out and drawn by Radoszkowski, — presumably one of the reasons why the specimen has become lost or destroyed. It was not present in Radoszkowski’s collection of Hymenoptera in the Krakow museum (Popov, 1960), nor anywhere else. The descrip- tion is superficial, no information being given on the mouthparts, antennae, etc.; but judging from the outline sketches of the scutellum and hind leg, there can be no doubt that Radoszkowski’s bee is the same species as the one described as glasunowi by Mora- witz, some years afterwards. This is evident also from the author’s remarks: “...... bords des 4me—Sme segments ventraux frangés des poils longs et couchés noirs, mélés de poils blancs, qui forment une espéce de brosse”. The illustrations of the sternal plates and Fig. 33—39. Paracrocisa spp., & structures; fig. 33—35, P. kuschakewiczi (Radoszk.), from Af- ghanistan, dorsal view of labrum (33), external view of right hind tibia and basitarsus (34), apex of same right hind tibia, more enlarged, interoventral view (35); fig. 36—38, P. sinaitica Alfken, from Gebel Elba, external view of right hind tibia and basitarsus (36), apex of same right hind tibia, more enlarged, interoventral view (37), and tergite 7, ventral view (38); fig. 39, P. guilochei (Dusmet), from Morocco, apex of right hind tibia, interoventral view (same magnification as fig. 35 and 37) genitalia, published by Radoszkowski in 1893, and obviously taken from the same specimen, are excellent and quite decisive. As neotype of P. kuschakewiczi I have selected a male in perfect condition from Morawitz’ collection in the Leningrad museum, which came as nearly as practicable from the original type-locality. It tallies the original description of kuschakewiczi, at the same time agreeing in all respects with authentic syntypic specimens of the same sex identified by Morawitz as P. glasunowi. The original description of male and female glasunowi is appropriate and very full, 304 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 the characters mentioned being applicable also to the other species, except that the author failed to mention the characteristic thick bristles covering most of the exposed surface of the last four gastral sternites of the male. The species exhibits but little variation, except in size. The ground colour of head, thorax and part of the legs in the majority of specimens is black, whereas the scutellum, femora and abdomen are definitely dark brown. All individuals appear to be quite mature. In Friese’s unlocalized male, the Konya female from Turkey, and the two large males from Iran and Afghanistan, the body is entirely black. Males are more profusely white-haired than females, in which all markings are more sharply outlined. Body shape and pattern of the male are practically as shown in the photograph (Pl. 2 Fig. 7—8) a Fig. 40—41. Paracrocisa kuschakewiczi (Radoszk.), & from Afghanistan; fig. 40, sternites 8 and 7, external view; fig. 41, right half of genital capsule, ventral (left) and dorsal view. Both halves of sternite 7 torn apart lengthwise and drawn separately for P. guilochei, the small size of the white spot on the first gastral segment of male kuschakewiczi (Pl. 2 Fig. 10) being rather unusual for that species. The brushy hairs on the abdominal sternites are mainly dark brown or black, except laterally at 3 (or 3—4), on which they are at least partly white. For more details, see Morawitz’ descrip- tions. Size variable. Total length: $ 13.0—15.5 mm, fore wing 9.0—12.0 mm; 9 12.0— 16.0 mm and 9.3—11.0 mm, respectively. Distribution. — Transcaspian region as far as Alma Ata, and into N. Iran. M. A. LIEFTINCK: Old World Melectine bees 305 Paracrocisa sinaitica Alfken (Pl. 2 Fig. 9; Fig. 36—38, 42— 43) Paracrocisa sinaitica Alfken, 1937, Konowia 16 : 173—175 (49 Wadi Ab-Orta, Sinai). — Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR 21 : 329 (not seen). Type material. — Egypt: Sinai Peninsula: 1 & (lectotype P. sinaitica Alfken, by present selection), SO. Sinai, Kneucker (written), Allotypus (print on red), Paracrocisa sinaitica m., det. J. D. Alfken 1937 (MNB). Further material. — Egypt: 1 g, Egypt, Wadi Digla, 29-31.5.1924, Coll. C.B.W. & T.W.K., Min. Agric., Slg. Alfken (ML); 1 8 (diss, fig. 36—39, 42—43 & Pl. 2 Fig. 9), SE Egypt, Gebel Elba, A. 8.3.38, H. Priesner (CP); 1 Q, Egypt, Wadi Hussein, 31.5.1919, coll. Adair, Min. Agric. (MNB). — Israel: 1 ©, Palestina, Ejn Geddi, 30.3, H. Bytinski-Salz (CBS). This conspicuous bee is sufficiently characterized in the generic diagnosis and key to the species. See, however, under P. gwilochei (Dusmet). The lectotype is a worn specimen that has lost most of its white thorax pubescence. A few additional remarks, taken from fresh examples, may suffice to recognize it. Male. — Characters of P. guilochei, but differing at first glance by the greater extent of white markings on all parts of the body. Labrum with the short and slightly raised hairs sparsely distributed, but all white; spot at base of mandibles also more conspicuous. Clypeus and summit of head all white, with narrow, transverse hairless spaces only at sides of lateral ocelli; lower portions of head mixed black and white. Mesothoracic collar more extensive, ill-limited posteriorly, reaching to beyond anterior margin of tegulae, which themselves also are partly white-haired. Knees of femora and outer faces of fore and mid tibiae white from end to end, those of hind tibiae on basal half only, their outer limits oblique, for the rest almost hairless, shining, and sculptured as in gwilochei; all tarsal segments covered with minute white hairs on a dark ground. Abdominal pattern similar to guilochei, but white patches larger and the one at sides of 1 extending further inward at base (Pl. 2 Fig. 9). Ventral surface of abdomen black, except small white patches at sides of 3, or 3—5. Apex of tergite 7 usually shallowly excised, with the rounded angles a little swollen and upturned (Fig. 38). Sternites 7 and 8, and copulatory apparatus, as in Fig. 42 and 43. Female. — Structurally quite similar to gwilochei, except that the dorsal process at the apex of the hind basitarsus is not spoon-shaped but triangularly pointed. The only avialable individual, from Wadi Hussein, is much darker than the male and may not even be conspecific. White marks on head reduced to two thick facial bars, one at each side, extending along margin of compound eyes below and above implantation of antennae, the latter being placed at about midway their lengths; remainder of clypeus and dorsal surface of head black. An additional, circular, dorsolateral white spot is placed at each side behind the eyes. White spots on thorax also smaller than in gzilochei, like those at base of all tibiae, which are completely isolated, occupying only one-fifth or less of the total external area. Gaster dull, finely black-haired above and underneath. White pubescent spots much smaller than in male and restricted to the sides, the pair on tergite 1 extending from base to apex but shaped like an inverted comma and barely visible from above; spots on 2—4 only little larger, those on 2 and 4 roundish, on 3 306 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 42—43. Paracrocisa sinaitica Alfken, & from Gebel Elba; fig. 42, sternites 8 and 7, external view; fig. 43, right half of genital capsule, ventral (left) and dorsal view. Both halves of sternite 7 torn apart lengthwise and drawn separately transverse and constricted at middle, all of them much smaller than the distance sep- arating them on dorsum. Apical border of sternites 1—4 rather broadly, of 5 more narrowly, impunctate, microscopically tessellate and shining, more broadly so than in guilochei. Sparse fringe of black subapical bristles as in that species. Pygidial plate not exposed. Total length: $ 15.0—16.0 mm, fore wing 11.0 mm; 9 14.0 mm (approx.) and 11.2 mm, respectively. Paracrocisa guilochei (Dusmet) (Pl. 2 Fig. 7—8; Fig. 39, 44—45) Melecta (Pseudomelecta) Guilochei Dusmet Y Alonso, 1915a, Mem. Real Soc. Esp. Hist. Nat. 8 (7a) : 331—333 (4 Tigui, Marruecos). Dusmet Y Alonso, 1915b, Bol. Real Soc. Esp. Hist. Nat. 15 : 257 (2 Imi-n-Tanut, Atlas, Marruecos; Gwilochei, supposition). Paracrocisa guillochei (sic), Popov, 1955, Trudy Zool. Inst. Akad. Nauk USSR, 21 : 330 (not seen). ? Melecta transitoria Pérez, 1896, Esp. nouv. Mellif. Barbarie, Bordeaux : 26 (& sine loc., rect. Djebel Bounda, Tunisie). Type material. — Morocco:1 ¢ (holotype M. guilochei Dusmet, PI. 2 Fig. 7), with label Tigui (written), Pseudomelecta Guilochei Dusm. & (Typo) Da la colecciön Escalada (three labels in Dusmet’s writing) (IEM). Further material. — Morocco: 1 & (diss, Pl. 2 Fig. 8 & Fig. 39, 44—45), environs of Melilla, 6.VI.1955, N. F. de Andrade (MZUC); 1 2, Maroc, Maharidja, Museum Paris, coll. J. de Gaulle 1919, with Melecta grandis (unknown hand) (MP); 1 2, Morocco: nr. Ouarzazate, 16.1V.1968, J. G. Rozen & E. Suissa coll. (AMNH). M. A. LIEFTINCK : Old World Melectine bees 307 One of the two new Melectini described by Dusmet in his “Apidos de Marruecos”, viz. Melecta Novellai Dusmet, is conspecific with M. albovaria (Erichs.), which is probably best considered a subspecies of M. punctata (F.). The second, also unique, is his M. (Pseudomelecta) Guilochei, now before me. With regard to body texture and nature of pubescence, the male of this insect approaches P. sinaitica Alfken so closely, that I first considered the differences between them (including those found in the copulatory apparatus) to be of little taxonomic importance. With further material of both coming to hand, this may indeed prove to be so. I am, however, unable to express any opinion about the amount of infraspecific variation that may be manifest in the various parts of the genital organs and other so-called primary sexual characters of these rare bees. No single Paracrocisa has yet been found in the coastal provinces between Morocco in the west, and Egypt and Israel in the east, so that possible intermediate forms are lacking. Consequently, I prefer for the present to keep these two species apart. In case of their unison, the name guilochei has, of course, priority over sinaitica. A still earlier described species is the enigmatic Melecta transitoria Pérez, based on a unique specimen from Tunisia, which could not, however, be recovered in J. Pérez’ collection at the Paris museum, and has probably become lost. Its description runs as follows: “07. M. transitoria. — 3. 11—12 mm. La tête, l'avant et le dessous du corselet couverts de forts poils blancs; 2 taches devant les appendices de l’écusson unies par une traînée latérale à la grande bande antérieure. Dessins de l'abdomen comme chez les Crocisa, c'est-à-dire en chevron aux deux lers segments; deux taches longues, sinuées, aux trois suivants; au 6° une tache ovale. Écusson très prolongé, émettant deux longues épines. Antennes longues, le 3° article deux fois plus long que large, les suivants une fois et demie. Tibias postérieurs tuméfiés, grossièrement tuberculés, portant au bout un éperon analogue à celui de la funeraria, plus long et plus aigu; le prototarse élargi, courbé et creusé. Les deux lers segments ventraux à peu près nus et très luisants, les suivants recouverts d'un épais duvet noirätre, plus abondant aux bords, frisé et relevé au bout, comme chez les Epeolus” — Dr. S. Kelner-Pillault kindly informed me in a letter, that there is a manuscript note in J. Pérez’ unpublished catalogue referring to this insect. It says: “4 juin, Djebel Bounda, Tunisie. Dessin de l’abdomen d'une Crocisa, villosité du thorax en partie, écusson tendant aussi à la forme crocisienne”. As follows from the above description, the species may well be identical with guilochei, and if so, the name transitoria would take precedence of all others, in which case it should be accepted as the proper name for the type species of Paracrocisa. The type of guilochei, still in good condition, agrees in almost all respects with the only other available specimen of that sex, both photographed on PI. 2 Fig. 7 and 8. Male. — Labrum as in Fig. 33, of P. kuschakewiczi, sides with thin apical tuft of raised white hair, the anterior border fringed with black, for the rest almost bare. No white at mandible bases. Antennal segment 3 in both dorsal and ventral view slightly but distinctly longer than 4, which equals 5 in length, the following segments being a little shorter. Clypeus for the greater part hairless (probably entirely white in fresh examples). Raised, finely branched hairs on top of head and mesothoracic collar shorter and less numerous than in P. sinaitica. Scutellar processes very short, only one-fourth (or less) length of main body, spike-like and directed straight back. Fore femora with 308 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 white posterior fringe, hairs about equal in length to diameter of femur; mid and hind femora covered sparsely with vestigial dark hairs. White appressed patches on outer faces of all tibiae ill-defined distad and occupying basal two-fifths only. Tarsal segments also sparsely white externally, the short hair on inner faces blackish. Ventral pubescence of abdomen, the sternal fringes included, black, except small tufts of white at sides of sternites 3 and 4. The density of the punctation on the thoracic dorsum and scutellum is obviously unreliable as a specific character, for in the type the mesonotal punctures are practically fused together, whereas in the second male the distance separating them, though much less than one puncture width, is smooth and slightly shining. Tergite 7 shaped similarly to that of P. sinaitica (Fig. 38), apex shallowly emarginate. Sternal plates 7 and 8 as in Fig. 44. Apart from the fact that the gonostylus, as compared with that of sinaitica, is more slenderly pointed, the pubescent flap to the inside of the latter also differs somewhat in size and shape. In ventral view, the strongly setiferous ventrobasal process appears to be small and tubercular; seen from within, however, it runs dorsad into the capsule as an elongate ridge at right angles to the main axis of the body. In dorsal view, the dorsobasal process is considerably longer than in kuschakewiczi, rod- or finger-like, resembling the shape it has in sinaitica. The remaining interior appendages (i.e., penis valves and spatha) are distorted in the dissected specimen and therefore useless for comparison with the other species (Fig. 45). Female. — Very similar to the male. Disk of labrum with few black bristles, lateral and apical fringes likewise black. Maxillary palpus slender, segments 1 and 6 of equal length, each about two-thirds as long as the remaining segments. Front of antennal scape with short, scanty appressed hairs; 3 hardly longer than 4, length and breadth ratio 3 : 2, succeeding segments only a trifle shorter. Anterior surface of head, with the Fig. 44—45. Paracrocisa guilochei (Dusmet), & from Morocco; fig. 44, sternites 8 and 7, external view; fig. 45, right half of genital capsule, ventral (left) and dorsal view (penis valves and spatha distorted and shown in displaced position) M. A. LIEFTINCK : Old World Melectine bees 309 exception of bare distal half of clypeus, white, as also a transverse band of longer hairs on vertex (subinterrupted by black in the median line); lower half of genal area and all ventral parts of head, black-haired. Pro- and mesonotum with the usual series of isolated white spots, the hypoepimeral area and upper half of mesepisternum clothed with long white hairs forming a large oval patch; sides for the rest black-haired. Shape, sculpture and pubescence of scutellum, as well as posterior thoracic segments, as in male. Thorax, coxae, femora and abdomen ventrally, black, sparsely black-haired. Outer faces of all tibiae with sharply defined white hair-spot occupying approximately their basal one-third; fore and mid tibiae beyond this finely tessellate-punctate, scantily bristled and strewn with thick spicules towards the end; dorso-apical process at mid tibia long and cylindrical. Hind tibia beyond hair patch smooth and shining, but surface uneven and clad with numerous short, thick spicules on raised tubercles, present also at the broadly rounded hind margin, beyond the subapical impressed area; inner spur only slightly undulated. All tarsal segments black-haired; hind basitarsus almost straight and parallel- sided, ending in a short, spoon-shaped external lobe. Total length: $ 14.0—15.0 mm, fore wing 10.0—11.0 mm; @ 14.0 mm and 10.5—11.0 mm, respectively. According to Dusmet (1915b), the females he received from the Atlas Mts. in Morocco (probably also in the Madrid museum), had lost too much of their freshness to allow of a proper identification. However, since the author compared them with the description of P. kuschakewiczi, pointing out also that the proportional lengths of the antennal segments differed from those of the type male of his own species, there can be little doubt that Dusmet’s examples were correctly assigned to P. guilochei. Afromelecta gen. nov. General appearance, Pl. 2 Fig. 11 and 12. The principal characters of this very distinct genus are enumerated in the key and will be understood more clearly from the specific description and illustrations. Of the three taxa included in Afromelecta, unfortunately only the type species can be fully characterized, the structural features of the other two being still incompletely known. Type species: Crocisa fulvohirta Cameron, 1905. Afromelecta fulvohirta (Cameron) comb. nov. (Pl. 2 Fig. 11—12; Fig. 46—53) Crocisa fulvohirta Cameron, 1905, Trans. S. Afr. Phil. Soc. 15 (4) : 247 (@ Cape). — Friese, 1909, Bienen Afrikas, Jena : 301, 312 (orig. descr. quoted; not seen). Melecta niveipes Enderlein, 1906, Stett. Ent. Ztg. 67 : 287—289 (Capland 3 9; davon 2 ® im Berliner, 1 9 im Stett. Zoolog. Museum). — Friese, 1909, Bienen Afrikas, Jena : 300—301 (verbatim quotation from Enderlein). — Bischoff, 1923, Deutsch. Ent. Zeitschr. H. 5 : 601 (E. kobrowi Brauns a synonym of M. niveipes End.). Epeolus kobrowi Brauns, 1909, Verh. zool.-bot. Ges. Wien, 59 : 10—12 (9 4 Kapland). — Meade-Waldo, 1913, Ann. Mag. Nat. Hist. (8) 12 : 97 (probably a synonym of E. fulvohirtus (Cam.)). — Friese, 1909, Bienen Afrikas, Jena : 298 (verbatim quotation from Brauns). Epeolus fulvohirtus, Meade-Waldo, 1913, Ann. Mag. Nat. Hist. (8) 12: 97 (probably syno- nymous with E. Zobrowi Brauns). — Brauns, 1926, Zool. Jahrb. Abt. Syst. 52: 206—207 (M. niveipes End. and E. kobrowi are synonyms). Type material. — South Africa: Q, lectotype of M. niveipes Enderlein, by 310 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 present designation, labelled Capland (print on blue) 31248 / Type (orange), Melecta niveipes 9 1906, det. Dr. Enderlein (MNB). Further material. — South Africa: 1 @, Capland (print on blue), 31249 / Type (orange), Melecta niveipes 4 1906, det. Dr. Enderlein, with additional label Melecta baeri Rad., det. Friese 1904 (MNB). 2 9, Delareg, W. Transvaal, 1.1917, Dr. Brauns, one with “3 specimens Epeolus kobrowi Brauns TM” (H.N. Empey’s writing (TMJ). 1 9, Delareg, W. Transvaal, 1.1917 (Brauns’s writing), Dr. Brauns / Typus (print on orange), 220.61 (red), Epeolus fulvohirtus Cam. = kobrowi Brauns, det. Friese 1910, Melecta fulvohirta Cam., det. Friese 1909 (NRS). 1 9, Bothaville, Orange Fr. St. 25.2.99, Dr. Brauns, Epeolus n. sp., det. Kohl (TMJ). 1 & (genit. diss. by H. N. Empey), 1 9, Florida Hills, 12 mi. from Johannesburg, Transvaal S. Afr., 31.XII.66, H. N. Empey, Epeolus kobrowi Brauns, compared with & and 9 types Transvaal Mus., det. Empey 1970 (ML). 1 9, Queenstown, Cape, S. Africa, 16.1.1965, H. N. Empey, Epeolus kobrowi Br., var., det. Empey (ML). 1 ©, erroneously labelled v. d. Wulp China ? (ML). 2 & (one diss., Pl. 2 Fig. 12 & Fig. 51—52), 1 ©, Annshaw {2}, S. Africa, Barrett 97—92, sub Melecta niveipes (BM). 1 9, Africa S. (BM). 1 9, Cape G.H. (bold capital print), Smith coll., pres. by Mrs. Farren-White 99—303 (BM). 1 ®, Natal, Weenen, 1.1926, H. P. Thomasset, pres. by Imp. Bur. Ent. 1929—407 (BM). The type of Cameron’s Crocisa fulvohirta, in the British Museum (Nat. Hist.) col- lection, is a discoloured specimen in poor condition. I have not examined this bee, but Meade-Waldo has, who transferred it to Epeolus and supposed it to be the same species as E. kobrowi Brauns. Some years previously, Friese had already suggested that Melecta niveipes Enderlein should also belong here, but Brauns (1926) was the first to establish the full specific synonymy of this remarkable South African bee. When Bischoff (1923 : 601, note) writes: “Epeolus kobrowi Brauns ist nach Vergleich der ausführlichen Be- schreibung mit den typischen Exemplaren der Melecta niveipes Enderlein zweifellos ein Synonym dieser”, it is obvious that he was inclined to place it near Melecta. In spite of that, Brauns still called it Epeolus fulvohirtus (Cameron)! It is of some interest to quote some of Brauns’ remarks on this insect which are included in his article: “Ich kann nun mitteilen, dass die drei Arten tatsächlich zu einer und derselben Species ge- hören, und muss nunmehr dieselbe Epeolus fulvohirtus heissen, obwohl der Name völlig unpassend ist, da die Behaarung schneeweiss ist, wie aus Enderlein’s und meiner Be- schreibung zu entnehmen ist. Cameron hat jedenfalls ein schmutziges oder verfärbtes Exemplar vor sich gehabt; seine Beschreibung ist daher auch fehlerhaft. Die Art ist für einen Epeolus sehr gross und aberrant, und kann leicht für eine Melecta angesehen werden, deren Habitus sie hat. Sie kommt im Norden von Südafrika, im Orange-Freistaat und Transvaal nicht selten vor und ist ein Parasit der grossen Anthophora (Habropoda) festiva Dours. Letztere ist dort eine sehr häufige Biene, findet sich aber auch im Süden der Kapprovinz.” (loc. cit.: 206—207). In the British Museum collection the specimens stood over the drawer label Melecta niveipes Enderlein, but none of them bear iden- tification labels. The lectotype 9 and first described & (or allotype) of the latter, in the Berlin Museum, agree in every respect with those in the British Museum and examples in other collections referred to above. E. kobrowi was described after 3 9 and 2 @ taken from December to February in Orangia (Bothaville and Reddersburg) and during January in the Cape Colony (Queenstown). According to Mr. H. N. Empey M. A. LIEFTINCK : Old World Melectine bees 311 Fig. 46—50. Afromelecta fulvohirta (Cameron), & and ® from S. Africa; fig. 46, 9 syntype E. kobrowi Brauns, from W. Transvaal, right side view of head; fig. 47, 2 from S. Africa, dorsal view of left mandible; fig. 48, Q from S. Africa, dorsal view of labrum; fig. 49, & from Annshaw, S.A., dorsal view of tergite 7; fig. 50, ® from the Cape, dorsal view of pygidial plate (in litt.), the types (9 and J') are in the collection of the Transvaal Museum (Jo- hannesburg) and compare very well with those taken by himself. The latter were gen- erously presented to me and will be deposited ultimately in the Leiden Museum. Brauns’ types, in the type cabinet, bear the nos. 740 (@) and 741 (g') and were caught by Brauns at Reddersburg (O.F.S.), 20.XII.1898 and 1.11.1899, respectively, bearing Brauns’ identification and type labels. This conspicuous bee resembles Paracrocisa most closely in general appearance and size. The body has the same elongate-cylindrical form, and although the abdominal segments are rather more arched, the gaster tapers to the end in the same way. Apart from the 312 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 primary sexual characters, male and female are practically alike, the thorax pattern being almost identical, differing in this respect from Melecta and Paracrocisa (and Pseudomelecta as well), in all of which the sexes are strikingly dimorphic. Male and female. — Inner mandibular projection (Fig. 47) slightly in advance of the middle, usually forming a small, oval, flattened tubercle which is occasionally obsolete and — like the apices of the mandible itself — worn off; long bristles are present around the tubercle, an apical tuft also at the carina to the outside of it, and a long fringe borders the mandible externally. Malar space distinct though linear. Labrum (Fig. 48) short (ratio of length and breadth 75 : 90 in male, 70 : 80 approx. in female); whole surface coarsely punctate, evenly clothed with conspicuous brush of long and stiff, suberect, dark reddish- to blackish-brown bristles, which are directed forward; apex variously produced, the tubercle on each side of the anterior emargination often less pronounced than in Fig. 48; subapical tubercle at middle of disk invariably present but often covered with dense hair. Clypeus broad, very little prominent, surface convex, sculptured with closely set, coarse and partly coalescent, punctures, rarely with weak median impunctate line; apical portion naked or with short, scanty, appressed hairs which become longer and more crowded together posteriorly until covering entire surface of parocular area; pile still longer, erect and feathery, behind antennae. Interantennal (frontal) carina in the Fig. 51—52. Afromelecta fulvohirta (Cameron), 6 from Annshaw, S.A.; fig. 51, sternites 8 and 7, external view; fig. 52, right half of genital capsule, ventral (left) and dorsal view form of a thick, distinctly raised carina, highest between antennae. Vertex and ocellar area bare, rather shining, finely tessellate, with few superficial punctures; postocular region and genal area clothed and fringed densely with raised white pubescence. Eyes large, widest well in advance of the middle and distinctly narrowed above; inner orbits little converging anterad and scarcely incurved on top; interocular distance almost or fully twice as long as basal width of labrum (ratio 100 : 55 in male, 100 : 50 in female), M. A. LIEFTINCK : Old World Melectine bees 313 interocellar and ocellocular distances about equal. Antenna normal, rather short and thick in both sexes; 3 about twice as long as its width at base and a trifle longer than 4 and next segments, which are only little longer than broad (ratio about 6 : 5). Thorax robust, punctation coarse and very dense, diameter of punctures about equal to their interspaces on disk of mesonotum, the latter with pair of small, oval, impunctate and shining areas, about half the size of tegulae, placed on each side upon middle somewhat in advance of tegulae, but usually barely showing through the pubescence. Scutellum in dorsal view more than four times broader than its median length, with pair of short, generally somewhat downcurved, posterolateral spines, which attain only half of the median length of scutellum or even less, the dorsal carina of each being distinct and the apices sharply acute. Pubescent pattern well defined (Pl. 2 Fig. 11—12), much as described for male Acanthomelecta, except that all white areas on dorsum of thorax (at least so in male) are more compact, composed of decumbent instead of raised hairs; anterior mesonotal spots either fused together, forming a broad collar of long, silky, decumbent hairs (male), or only with als and p/sa coalescent in front of tegulae, and with ms broader than in Acanthomelecta (female). Entire upper three-fourths of thoracic sides clothed with very long, decumbent, finely branched hairs forming a dense patch entirely overlying the integument and occupying also parts on each side of scutellum and lateral propodeal areas; for the rest the sides are black-haired with fringes of white at lateroventral border, in front of white tufts at the coxae. Legs quite normal, outer faces of femora in both sexes finely, rather closely punctate, more sparsely and superficially so inward; fore femur with very long and dense, mid femur with somewhat shorter posterior fringe of white, hind femur sparsely black and white only at ventral carina, all hairs on hinder pair much shorter than diameter of femur; outer faces of tibiae and tarsi white-haired, the tibiae of mid and hind legs covered with numerous short black spicules shining through the pubescence. No external pad of felt-like hair on mid tibia of male; hind tibiae of male with short, sharp, dorso- apical longitudinal crest which curves around its distal border, this apical carina lacking in female. Male hind basitarsus very slender, laterally compressed, almost parallel-sided after the basal constriction, nearly five times as long as its greatest width and distinctly outbent, its outer face clothed with short appressed hair; in female almost straight, not at all compressed or flattened, much narrower still than in male and clothed with bristly hair. Wings (Pl. 2 Fig. 12) long, fore wing membrane strongly smoky, the posterior and outer cell areas hyaline, as is also the hind wing except apically; marginal cell more than four times (4.1—4.5) as long as broad; length of pterostigma, marginal cell and distance from the latter to wing apex, in the ratio of 30 : 80.3 : 100. Abdomen with two first segments subequal in length, or first a trifle longer than second; black, all segments dull, finely superficially punctate, punctures smaller than interspaces on basal portions but crowded together posteriorly and extending to near posterior border of segments, above as well as underneath. Pubescence very short, black, with sparse fringe of longer subapical bristles on sternal surfaces. White dorsal marks similar in both sexes, as shown in Pl. 2 Fig. 11—12; sternites clothed sparsely with short black hair, 2—5 (male) or 2—4 (female) each with large, pure white pubescent spot placed transversely on either side. Tergite 7 of male shaped as in Fig. 49, its surface clothed densely with long, appressed, blackish brown hairs on either side of a somewhat spatulate median area, the apical portion of which is bare, flattened and finely tessellate, 314 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Fig. 53. Afromelecta fulvohirta (Cameron), & from Annshaw, S.A., right gonostylus with append- ages, dissected out and viewed from within; ventrobasal process removed and not shown. Fig. 54. Acanthomelecta bicuspis (Stadelmann), & holotype Melecta spinosa Friese from Usambara, right gonostylus with appendages, dissected out and viewed from within; dbg = dorsal branch of gonostylus; vbg = ventral branch of gonostylus; GONS = main body of gonostylus; dbp = dorsobasal process; vbp = ventrobasal process (omitted in fig. 53) with slightly swollen and ridge-like margin. Posterior margin of sternal plates 2—5 almost straight; sternite 6 of male protuberant, but apex rounded, surface with very slight median impression, clothed densely with fine, dark brown decumbent hair. Sternites 7 and 8 shaped as in Fig. 51. Genital capsule of large size (2.3 mm, incl. gonostylus), of very complex structure. General appearance in ventral and dorsal aspect, as in Fig. 52. After removal of the ventrobasal process of the right half, the appendages of the gonocoxite, viewed from within, are as shown in Fig. 53. The ventrobasal process of the right half is shown only in Fig. 52; it is folded lengthwise and consists of a heavily sclerotised basodorsal portion and a large, horizontally placed, thin, semitransparent ventral membrane. The dorsal (inner) portion is heavily sclerotised, directed into the capsule as a lanceolate rod-like apodeme; the rounded outer portion of the membrane is also firm and beset with longish hairs, its thin distal (inner) part gently incurved in ventral view, sub- truncated apically and bordered mesad by a broad, parallel-sided, subhyaline marginal strip, which is transversely striated (Fig. 52). The dorsobasal process of the gonocoxite is detached from the gonostylus and overlaps most of the latter in dorsal view; it consists of two separated parts, which are fused together only at extreme base: one small out- wardly convex subcircular dorsal lobe covered with very long, strong setae (dbp), and a much longer, more laterally placed, broadly spatulate incurved process (dhg) whose entire inner surface is clothed densely with similar but still longer setae, the latter almost concealing the underlying gonostylus (Fig. 52). Gonostylus likewise incurved, rather finger-like and tapering rapidly to a bluntly pointed tip; its basal part above is fringed M. A. LIEFTINCK: Old World Melectine bees 315 rather densely with long setae, the apex bearing only short hair (Fig. 53). Pygidial plate of female occupying about median one-third of tergal dorsum, shaped as in Fig. 50; surface flat or almost so, somewhat shining, finely reticulate; colour dark chestnut brown with slightly raised and almost black side margins. Total length: g 14.0—15.0 mm, fore wing 11.5—12.0 mm; 9 13.0—16.0 mm and 11.0—12.0 mm, respectively. Host-relationship. — With reference to Brauns’ observation, cited above, on the dis- tribution of this species, and its simultaneous occurrence with Anthophora (Habropoda) festiva Dours 1), in parts of South Africa, it is perhaps worthy of note that there are several other anthophorines which may be looked upon as foster bees of Afromelecta. Besides H. festiva Dours, labelled as from British E. Africa and the Cape Province, the collection of the British Museum (Nat. Hist.), for instance, includes species such as H. determinata Friese, from Nyasa (Tanzania) and Portuguese E. Africa, and H. bettoni Ckll., reportedly from Abyssinia and British E. Africa. The male of H. bettoni from Ethiopia is quite extraordinary in that the basitarsus of the fore leg is furnished with a very conspicuous beard-like tuft of hairs, while others show unusually specialised antennal and leg structures. As far as I have been able to ascertain, the basic characters of this African section do not coincide with those of Mediterranean and Eurasian Habropoda s. str., as redefined by me recently (Lieftinck, 1966), the former being not even homo- geneous in itself. So far, I have studied only the copulatory apparatus of a male from Tanganyika, in the Leiden Museum, which apparently belongs to an undescribed species of the same section. The apical sternal plates and genitalia of this bee are altogether different from those of true Habropoda, so that it will probably be necessary to erect a new genus for the reception of this and allied African species of the group. Afromelecta spec. indet. Material. — Ethiopia: 1 9, with printed label: Abyssinia, Higo Samula, 30. 10. 11, R. J. Stordy, 1912—329 (BM). A specimen in perfect condition, evidently a close ally of A. fulvohirta (Cam.) and closely resembling it, but distinguished by having longer scutellar spines, which in side view are equal in length to the main body of the lobes; these spines are cylindrical at their bases, tapering to a point instead of being laterally compressed and strongly carinated above. The specimen differs further in that (1) the first gastral tergite is slightly less closely punctate; (2) almost the entire lower half of the thoracic sides is black- instead of white-haired; and (3) the apical one-fifth of the mid tibiae and apical one-half of the hind tibiae are black-haired on their outer faces, whereas in fulvohirta the tibiae are entirely white except at the tips, which are narrowly bordered with black. With the discovery of the male and more material of both sexes from the same region, this may prove specifically distinct from A. fulvohirta (Cam.). Afromelecta spec. indet. Material. —Kenya:1 g (dissected, but terminalia all missing), with three printed 1) Cameron (1905: 255—256) described a second large-sized bee, Habropoda capensis Cam., measuring 17 mm in length, from the Cape Colony (Bivak Kloof) and Transvaal. This is very likely the same species as festiva Dours. 316 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 labels: Brit. C. Africa, Kuja Valley, S. Kavirondo, 4000 ft., Apr. 30—May 1, 1911, S. A. Neave, 1912—193 (BM). A more slenderly built insect than A. f#lvohirta, immediately distinguished from it by the slightly shorter Jabrum, the surface of which is not clothed with a dense brush of brown hairs but with thinner and pure white pubescence. Also, the scutellum is less coarsely punctate and the spines are longer, slenderer, with more sharply pointed tips, than in fulvohirta. Further differences are found in the abdomen, which is narrower and distinctly more slender, the tergites are less closely punctate, with broader impunctate hind margins and a more shining surface; lastly, the hind basitarsus is narrower and more definitely hollowed out externally. Although agreeing with A. fulvohirta in many respects, this is undoubtedly a different species. However, by the absence of the apical segments and genital organs, I prefer to leave it without a name until more material will become available. Acanthomelecta gen. nov. A monotypic genus of medium-sized bees, with melectine characters intermediate between those of Afromelecta, Melecta and Paracrocisa. In the generic key and the following descriptions I have endeavoured to characterize male and female as complete as possible. Although it should be borne in mind that these features are taken from a single individual of either sex, it was found impossible to fit this remarkable species into any genus so far recognized. This necessitates the erection of a separate taxon to hold it. Acanthomelecta is distinguished from Melecta by the greater length of the first gastral tergite, the absence of a well-developed tarsal arolia, the totally different shape of the scutellum, with its extraordinary spines, and the much shorter vestiture covering the head and thoracic segments. By the form of the head and apical sternal plates, the male of Acanthomelecta approaches Afromelecta most closely, but the scutellar spines here are short and the genitalia of the males are altogether dissimilar; moreover, the maxillary palpi in the three species presently attributed to Afromelecta are reduced to one or two short segments, whereas Acanthomelecta is probably more primitive by having a 6-seg- mented palpus. In spite of this, it must be admitted that the two genera appear to be more nearly related to one another than either of the two can be linked with Paracrocisa, which comprises a compact group of closely similar forms, distinguished — among other characters — by the characteristic structure of the antennae, legs and genital organs of the male. It would be of great interest to examine additional material from the typical localities of A. bicuspis and to discover more species from other parts in Africa, in order to find out whether the most significant characters separating the former from Afro- melecta are constant or not. It is by no means certain that they are; and if, for example, specimens would occur which possess maxillary palpi having a varying number of seg- ments or showing somewhat intermediate forms of scutella, then Acanthomelecta should be merged in Afromelecta. As this seems unlikely, I assume for the present that more species do exist which will prove to fit into either of the two genera here recognized. Nothing, of course, can be said about the host-relationship of this bee. Like Afro- melecta, it may be parasitic on the Habropoda-like bees belonging to the assemblage typified by H. festiva Dours. This is a purely African group, practically unstudied yet, and certainly not congeneric with Habropoda F. Smith, as at present interpreted (see Lieftinck, 1966). Type species: Crocisa bicuspis Stadelmann, 1898. M. A. LIEFTINCK : Old World Melectine bees 317 Acanthomelecta bicuspis (Stadelmann) comb. nov. (Fig. 54—57) Crocisa bicuspis Stadelmann, 1898, Hym. Ost-Afrika's in K. Mobius, Thierwelt Ost-Afrik., etc. 4 (3) : 28 (nom. nud.!), pl. fig. 6 (thorax dorsally) (Q Ujunjoni, Stuhlmann). Melecta bicuspis, Meyer, 1921, Archiv f. Naturgesch. 87 : 177 (not seen). Melecta spinosa Friese, 1922, Zool. Jahrb. Abt. Syst. 46 : 33 (& Usambara). Syn. nov. Type material. — Tanzania:1 Q (discoloured, right fore wing missing), Ujun- joni 24.9 / Stuhlmann (written), Crocisa n. sp. bicuspis (ditto), Croc. bicuspis Stadel- mann Type, Dr. Enderlein (Enderlein det.), Type (print on light red); holotype Crocisa bicuspis Stadelmann (MNB). — Tanzania:1 g (all wings clipped off and wanting ! diss., Fig. 56—57), D.O.-Africa, Usambara, 6. 1903 (print), Melecta spinosa Fr. det. Friese 1908 (Friese’s writing), Type (print on dark red); holotype Melecta spinosa Friese (MNB). It is surprising that Friese failed to acknowledge the existence of Möbius’ “Thierwelt Ostafrikas” (1898) in his own great work, thus neglecting to deal with the bees des- cribed — or, more properly, listed — by Stadelmann, whose collection could have been studied so easily at the Berlin museum. This did not happen, and so it can be explained why the types of Crocisa bicuspis and Melecta spinosa were never confronted and left unstudied until the present day. Although Stadelmann’s account is incomplete, ending abruptly on page 74, it contains a number of figures which, according to Art. 16(a) VII of the Code, validate several of the proposed nomina nuda, including the present one. A fairly large-sized melectine, superficially resembling Afromelecta and Paracrocisa, but differing from both in details of structure. The two specimens, one of either sex, are unique and unquestionably conspecific. Female (holotype C. bicuspis). — A soiled and discoloured specimen, due to wetting, but clotted pubescent pattern clearly discernible. Inner tooth at about halfway length of mandible reduced to a mere convexity. Max- illary palpus as in Fig. 55. Labrum squarish (length : breadth = 75 : 80), widest at middle, surface slightly concave, closely punctate, punctures lacking interspaces; anterior border a little produced, apex distinctly upturned and with small crescentic emargination between bluntly triangular projections; whole margin fringed with brown bristles; tomentum short and dense, decumbent, the dark brown hairs directed anterad, inter- spersed with few, much longer, erect bristles. Clypeus less protuberant than lower diameter of eye, surface slightly convex, dull, very finely, closely punctate, lacking inter- spaces, clothed with very short, appressed white pubescence, the hairs becoming longer posteriorly. Interantennal (frontal) carina low, texture and pubescence similar to that upon hinder part of clypeus. Paraocular area clothed also with appressed tomentum, the white hairs becoming long, tufty and intermixed with black behind antennae; vertex almost bare, surface somewhat shining, superficially, not very closely punctate, the punctures of two sizes. Postocular and genal areas with long raised hairs, longest and white behind eyes, black at middle. Eyes large, shaped as described for Afromelecta, but inner orbits con- verging anterad and slightly incurved above; interocular distance slightly less than twice as long as basal width of labrum (ratio 100 : 53); interocellar and ocellocular distances as in Afromelecta. Antenna normal, slenderer than in Afromelecta, but scape similar; segment 3 more than two times longer than broad at base and subequal to next segments, which are only little longer than broad (ratio about 6 : 5). Thorax robust, whole surface 318 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 including scutellum and tegulae, dull, coarsely punctate, the punctation denser than in Afromelecta; scatellum in dorsal view more than four times broader than its median length, armed at either side of convex and strongly declivous median part with a long, robust, laterally compressed and carinated spur, which exceeds the median length of scutellum and is directed almost straight back; these spurs are more widely apart than the processes of Afromelecta and almost twice as long as they are in fulvohirta, the slender tips being acutely pointed. Pubescent pattern identical to that of the male, described below. Legs normal, all parts finely punctate, clothed with short pubescence, long pure white fringes only at ventral carinae of fore and mid femora; pile at outer faces of mid and hind tibiae decumbent, pure white along full length of fore tibia, and Fig. 55. Acanthomelecta bicuspis (Stadelmann), 9 holotype from Ujunjoni, maxillary palpus; scale- line = 0.5 mm. Fig. 56—57. Acanthomelecta bicuspis (Stadelmann), & holotype Melecta spinosa Friese from Usambara; fig. 56, sternite 8 and 7, external view, scale-line 1.0 mm; fig. 57, right half of genital capsule, ventral (left) and dorsal view, scale-line 1.0 mm about basal one-third of mid and hind tibiae, darker (discoloured ?) on tarsi. Mid and hind tibiae moreover covered with short, but very strong, suberect setae, especially so on hinder pair. Hind basitarsus shaped much as in female Afromelecta. Hind tibial spurs unequal, slightly curved, brown; tarsi as described in the key. Wings yellowish brown, much darker towards apex of radial cell and in apical part of fore wing. First sub- marginal on fore wing not much longer than diameter of third; marginal cell long, about three times (3.2) as long as broad; length of pterostigma, marginal cell and distance from the latter to wing tip in the ratio of 20.7 : 108 : 100; nervulus postfurcal for almost twice its own length. Abdomen elongate, tapering gradually posteriorly; gastral segment 1 slightly longer than 2. All tergites moderately shining, punctation much finer than on thorax, the punctures evenly distributed almost as far as posterior margin of segments and smaller than interspaces; punctation of sternites similar. Pygidial plate almost flat, rather broadly M. A. LIEFTINCK : Old World Melectine bees 319 triangular, a little less than twice as long as its width at base, side margins slightly raised, apex subobtuse; surface even, finely tessellate-punctate. Pubescence short, de- cumbent, black; tergites 1—4 with sharply defined white patches and spots composed only of finely branched appressed hairs: a large subtriangular patch, pointing inward, on each side of 1, and elongate paired transverse spots on 2—4, successively smaller posteriorly, all of them separated by black at middle by a distance greater than their own diameter. Sternites black-haired (discoloured ?). Total length: 14.0 mm (approx.), fore wing 10.3 mm. Male (holotype M. spinosa). — The unique specimen, though lacking its wings, is in a good state and agrees with the description, which requires the following corrections and emendations. Mouth-parts almost exactly as in the female: inner projection of mandibles a low crescentic convexity. Whole surface of labrum covered with dense, decumbent and raised, bristly hair, the short pile composed of white and dark brown tomentum, the hairs growing longer and almost black upon distal one-third. Pile on clypeus extremely short and appressed, silvery-white, much denser and longer upward than in front. Shape and pubescence of head, inclusive of the antennae, otherwise exactly as in the female (i.e., antenna not at all reaching scutellum and 2nd flagellar segment not longer than 3rd, as stated erroneously by Friese !). Mesonotum, scutellum and posterior thoracic segments closely, coarsely and rather deeply punctate; punctures on mesonotum at least as wide as the interspaces, more irregular and partly confluent on scutellum (including its processes basally) and posterior areas; upper surface somewhat shining and showing through the rather long, black pubescence, except where this is replaced by white patches of still longer, erect and feathery hairs, which form a sharply defined Thyreus-like pattern (apparently quite similar to that of the female), as follows: transverse paired a/s and longitudinal ms forming together an almost T-shaped anterior mark; large, isolated, subcircular p/sa and pls, the latter most conspicuous; and an equally large, subtriangular, median patch of white upon the vertical face of the scutellum, consisting of more decumbent, radiating plumose hairs, which are directed upward and forward. Sides and posterior areas of thoracic segments marked similarly to Afromelecta. Legs less slender, femora and tibiae somewhat more swollen and tarsi a little shorter, than in female; otherwise similar, lacking modifications; outer faces dull, coarsely closely punctate, inner faces more shining, sparsely and finely punctate. Pubescence as in female; long white fringes at base and at femora; short, appressed, black mixed with white, on following parts: whole outer face of fore tibiae and tarsi, basal three-sevenths of mid and hind tibiae, with narrow fringe at extreme tip, and outer faces of all tarsal segments. No external pad of felt-like hair on mid tibia. Hind tibia without any indication of a dorso-apical longitudinal carina. Hind basitarsus laterally compressed, straight and slender, slightly broadened from base to apex, almost five times as long as its greatest width near apex, surface scarcely con- cave externally. Gastral tergite 1 distinctly longer than 2; all abdominal segments black, somewhat shining, with slight oily reflections. Punctation fine, superficial, the punctures evenly distributed, much smaller than interspaces and extending to near posterior margin of seg- ments, above as well as underneath. Pubescence very short, black, with sparse fringe of longer subapical bristles on sternal surfaces, there being no dense subapical comb of strong bristles ventrally. A row of snowy-white lateral marks, progressively smaller from before 320 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 backward, on tergites 1—6; patches on 1 shaped as in female, though larger, subrectangu- lar, with convergent inner border, the intero-apical angles of the spots pointing inward; distance separating these spots posteriorly equal to their own diameter; all hairs finely branched and decumbent, except small tufts of raised hair laterally at extreme base. Spots on succeeding segments transverse, elongate-oval. Sternites clothed sparsely with short black hair, 2—5 each with distinct, pure white pubescent spot placed transversely at either side. Tergite 7 shaped much as in Afromelecta fulvohirta (Fig. 49), but finely tessellated, naked, midapical portion more abruptly narrowed basad and apex squarely cut off, with upturned, shallowly excised posterior margin. Sternite 6 protuberant, but apex rounded; surface very slightly impressed at middle, clothed densely with dark decumbent hair. Sternites 7 and 8 shaped as in Fig. 56. Genital capsule of large size (2.4 mm, incl. gonostylus). Ventrobasal process of gonostylus shaped similarly to Melecta, forming a small, heavily sclerotised, subrectangular and densely setiferous lobe projecting from the inside. Gonostylus broadest at base, rather strongly incurved and finger-like, setae on basal part above long and strong, surface for the rest clothed sparsely with finer and much shorter setae. 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Some Malaysian bees of the family Anthophoridae (Hym., Apoidea). — Treubia, hors sér.: 57—138, 79 figs., pl. 42. ——, 1959a. Notes on some eighteenth century bees of the genus Thyreus Panzer, with des- cription of a new species (Hym., Apoidea). — Tijdschr. v. Ent. 102 : 17—34, 20 figs., 2 pls. —. 1959b. Revision of the Indo-Australian species of the genus Thyreus Panzer (= Crocisa Jurine) (Hym., Anthophoridae). Part 2. Thyreus nitidulus (Fabricius). — Nova Guinea, new ser. 10 : 99—130, 50 figs., 2 pls. —, 1962. Revision of the Indo-Australian species of the genus Thyreus Panzer (= Crocisa Jurine) (Hym., Apoidea, Anthophoridae), Part 3. Oriental and Australian species. — Zool. Verhand. Leiden, 53 : 1—212, 72 figs., 3 pls. —, 1966. Notes on some Anthophorine bees, mainly from the Old World (Apoidea). — Tijdschr. v. Ent. 109 : 125—161, figs. 1—92. —, 1968. A review of Old World species of Thyreus Panzer (= Crocisa Jurine) (Hym., . Apoidea, Anthophoridae). Part 4.. Palearctic species. — Zool. Verhand. Leiden, 98 : 1—139, 38 figs., 4 pls. ———, 1969. The melectine genus Eupavlovskia Popov, 1955, with notes on its distribution and host relations (Hym., Apoidea, Anthophoridae). — Tijdschr. v. Ent. 112 : 101—122, figs. 1—8. Linsley, E. G. 1939. A revision of the Nearctic Melectinae (Hym., Anthophoridae). — Ann. Ent. Soc. Amer. 32 : 429 —468, 9 figs. — , 1943a. Notes on the habits of Melecta sierrae Linsley. — Pan-Pacific Ent. 19 : 160. ——, 1943b. The genus Melecta in Eastern North America and Porto Rico (Hym., Antho- phoridae). — J. New York Ent. Soc, 51 : 225—227. —, 1958. The ecology of solitary bees. — Hilgardia (Berkeley), 27 (19) : 543—599. figs. ———, & J. W. MacSwain, 1942. The parasites, predators, and inquiline associates of Anthophora linsleyi. — Amer. Midland Nat. 27 : 402—417, 11 figs. Meade-Waldo, G. 1913. Notes on the Apidae (Hym.) in the collection of the British Museum, with descriptions of new species. — Ann. Mag. Nat. Hist. (8) 12 : 92—103. Michener, C. D. 1948. Notes on the American bees of the genus Melecta (Hym., Apoidea). — Proc. Ent. Soc. Wash. 50 : 15—18. , 1953. Comparative morphological and systematic studies of bee larvae with a key to the families of hymenopterous larvae. — Sci. Bull. Univ. Kansas, 35 : 987—1102, 287 figs. 322 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Mitchell, T. B. 1962. Bees of the Eastern United States, 2. — Tech. Bul. 152, North Carol. Agric. Exper. Sta., 557 pp., 134 figs., 1 pl. Möczär, M. 1957. Méhfélék (Apidae) ir Fauna Hung. 13, Hym. 3, 76 pp., 22 fig. Morawitz, F. 1875. Hym. Mellifera I, in Fedchenko's Reise Turkestan, etc. Zool. (In Russian). — Mem. Imp. Anthrop. & Ethn. Moscou, 19 (2), 160 pp. , 1876. Hym. Mellifera I, in Fedchenko's Reise Turkestan, etc. Zool. — Mem. Imp. Anthrop. & Ethn. Moscou, 21 (3), tab. II figs. 26—27 (col. fig. 9 6 P. fuscipennis Mor.). , 1895. Beitrag zur Bienenfauna Turkmeniens. — Horae Soc. Ent. Ross. 29 : 1—76. Osychniuk, G. Z. 1970. Apoidea-Colletidae, ir Fauna Ukraine, 12 (4) (In Russian). — Inst. Zool. Akad. Nauk Ukrainskoi SSR, Naukova Dunka, Kiev. 158 pp., 77 figs. Pérez, J. 1883. Contribution 4 la Faune des Apiaires de France, 2. — Act. Soc. Linn. Bordeaux, 37 (4e sér., t. 7 live. 1) : 205—378, pls. 13—14. , 1896. Espéces nouvelles de Melliféres de Barbarie (Diagnoses préliminaires). — Bordeaux, 30 déc. 1895 : 1—64. Popov, V. B. 1950. The genus Amegilla Friese (Hym., Apoidea) (In Russian). — Entom. Oboz. Akad. Nauk USSR, 31 : 257—261, figs. ——, 1955. Generic classification of Palearctic Melectini (Hym., Anthophoridae) (In Russian). — Trudy Zool. Inst. Akad. Nauk USSR, 21 : 321—334, 4 fig. —, 1960. On Radoszkowski’s collection of Hymenoptera (In Russian). — Entom. Oboz. Akad. Nauk USSR, 39 : 237—240. Porter, J. C. 1951. Notes on the digger-bee Anthophora occidentalis and its inquilines. — Iowa St. Coll. J. Sci. 26 : 23—30. Radoszkowski, O. 1865. Tribu des Melectites. — Horae Soc. Ent. Ross. 3 : 53—60, tab. 1, figs. 1—3. , 1890. Etudes hyménoptérologiques. Descriptions d’espéces nouvelles de la faune Russe. — Horae Soc. Ent. Ross. 25 (1—2) : 244—248, pl. II. , 1893. Revue des armures copulatrices des males des genres: Crocisa Jur. ...... Acanthopus Klug. — Bull. Soc. Imp. Nat. Moscou, new ser. 7 (2—3) : 163—188, pls. IV—VII. Rayment, T. 1935. A Cluster of Bees. — Sydney, 752 pp., figs. & pls. Rozen, J. G. 1969a. The larvae of the Anthophoridae (Hym., Apoidea), Part 3. The Melectini, Ericrocini, and Rhathymini. — Amer. Mus. Novit. 2382 : 1—24, 56 figs. , 1969b. The biology and description of a new species of African Thyreus, with life history notes on two species of Anthophora (Hym., Anthophoridae). — J. New York Ent. Soc. 77 : 51—60, 11 figs. Semichon, L. 1906. Recherches morphologiques et biologiques sur quelques Melliféres solitaires. These, Fac. Sci. Paris : 281—442, pls. 13—15. , 1922. Sur la nymphe de Melecta armata Panzer (Hym. Apidae). — Bull. Soc. Ent. France, no. 14 : 192—194, 2 figs. , 1925. L'état larvaire de Melecta armata Panzer (Hym., Apidae). — Bull. Soc. Ent. France, no. 18 : 305—306. Stadelmann, 1898. Die Hymenopteren Ost-Afrika's, in K. Mobius, Die Thierwelt Ost-Afrikas und der Nachbargebiete, 4 (3) : 1—74, figs. Stoeckhert, F. K. 1954. Fauna Apoideorum Germaniae. — Abh. Bayer. Akad. Wiss. math.-naturw. KI., N.F. 65 : 5—87. Strand, E. 1913. H. Sauter’s Formosa-Ausbeute. Apidae I (Hym.). — Suppl. Entom. 2 : 23—67. Sustera, ©. 1958. Übersicht des Systems der paläarktischen und mitteleuropäischen Gattungen der Superfamilie Apoidea (Hym.) (In Czech). — Acta Ent. Mus. Nat. Pragae, 32 : 443—463 3 figs. Torchio, P. F. & N. N. Youssef. 1968. The biology of Anthophora (Micranthophora) flexipes and its cleptoparasite, Zacosmia maculata, including a description of the immature stages of the parasite (Hym., Apoidea, Anthophoridae). — J. Kansas Ent. Soc. 41 : 289—302. M. A. LIEFTINCK: Old World Melectine bees 323 ALPHABETICAL INDEX OF NAMES (Synonyms are in italics) abdominalis, Thyreus 258 abrupta, Anthophora 259 Acanthomelecta 256, 262 (key), 316 acervorum, Anthophora 256 aeneiventris, Anthophora 299 aestivalis, Anthophora 256 affinis, Thyreus 258 Afromelecta 256, 263 (key), 309 albigena, Amegilla 258, 259 albovaria, Melecta 307 amboinensis, Thyreus 259 Amegilla 258 andrewsi, Amegilla 258 Anthophora 255 Asaropoda 258 aspasius, Thyreus 258 atlantica, Melecta 256 atripes, Melecta 295 atripes, Pseudomelecta 257, 295 atroalba, Pseudomelecta 257, 286 (key), 293 baerii, Melecta 284 baerii, Pseudomelecta 282 bettoni, Habropoda 315 bicuspis, Acanthomelecta 256, 317 bicuspis, Crocisa 316, 317 bombiformis, Asaropoda 258 borealis, Anthophora 259 braunsiana, Anthophora 258 bucconis, Habropoda 270 buruensis, Amegilla 259 caeruleopunctatus, Thyreus 255, 258 caesareae, Melecta 299 caesareae, Paracrocisa 299 californica, Melectomorpha 255, 259 Callomelecta 267, 269 callura, Melecta 256 capensis, Habropoda 315 caroli, Amegilla 258 ceylonicus, Thyreus 258 chalybeia, Pseudomelecta 257, 258 (key), 296 cinerea, Emphoropsis 256 corpulenta, Melecta 284 corpulenta, Pseudomelecta 284 curta, Anthophora 259 cyrtandrae, Amegilla 258 determinata, Habropoda 315 diacantha, Melecta 287 diacantha, Pseudomelecta 257, 286 (key), 287 (key) edwardsii, Anthophora 257 edwardsii, Melecta 256, 259 Elaphropoda 257, 270, 281 elegans, Amegilla 258 elephas, Amegilla 258 Emphoropsis 256, 257, 281 Epeolus 307, 310 erratica, Elaphropoda 257, 282 Eupavlovskia 256, 263( key), 267 feronia, Amegilla 258 festiva, Habropoda 256, 310, 315 fimbriata, Amegilla 259 flammeozonata, Amegilla 259 flexipes, Anthophora 255, 259 formosana, Melecta 274 formosana, Protomelissa 257, 273 (key), 274 fulvitarsis, Anthophora 256 fulvohirta, Afromelecta 256, 309 fulvohirta, Crocisa 309 fulvohirtus, Epeolus 309 funeraria, Eupavlovskia 256, 267 fuscipennis, Melecta 290 fuscipennis, Pseudomelecta 257, 286 (key), 290 garrula, Amegilla 259 gemmatus, Thyreus 258 glasonovi, Melecta 302 glasunowi, Paracrocisa 302 guillochei, Paracrocisa 306 guillochei, Melecta 306 guilochei, Paracrocisa 257, 302 (key) Habropoda 257 habropodae, Callomelecta 281 habropodae, Protomelissa 257, 270 (key), 281 hanseni, Anthophora 257 himalayana, Melecta 273 himalayana, Protomelissa 257, 273 (key) himalayensis, Thyreus 258 histrionicus, Thyreus 258 impatiens, Elaphropoda 257 insidiosa, Callomelecta 282 insidiosa, Protomelissa 257, 272 (key) interrupta, Melectomorpha 259 irena, Thyreus 258 iridescens, Protomelissa 257, 272 (key), 277 irisanus, Thyreus 259 jacobi, Amegilla 258 kobrowi, Epeolus 309 kuchakevitzi, Paracrocisa 302 kuschakewiczi, Paracrocisa 257, 301 (key), 302 lamprosoma, Crocisa 258 lieftincki, Thyreus 258 linsleyi, Anthophora 255, 259 luctuosa, Melecta 255, 256 lugubris, Thyreus 258 maculata, Zacosmia 255, 259 Melecta 255, 256, 264 (key) 324 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 Melectomimus 257 Melectomorpha 259, 284 Melissa 269 Micranthophora 259 murrayensis, Amegilla 258 nereis, Thyreus 258 nitidulus, Thyreus 258 niveipes, Melecta 309 nivosa, Melecta 284 nivosa, Pseudomelecta 284 novaehollandiae, Thyreus 259 novellai, Melecta 307 obscura, Eupavlovskia 256, 267 orbatus, Thyreus 259 pacifica, Melecta 255, 256 Paracrocisa 257, 266 (key) Paramegilla 257 pendleburyi, Callomelecta 282 pendleburyi, Protomelissa 257, 272 (key) plagiata, Anthophora 256 plurinotata, Melecta 291 potanini, Anthophora 299 praestans, Thyreus 259 Protomelissa 257, 261 (key), 267 Pseudomelecta 257, 265 (key), 282 pulchra, Amegilla 255, 258 punctata, Melecta 256, 307 quadrifasciata, Amegilla 258 quadrimaculata, Anthophora 259 radoszkowskii, Habropoda 281 ramosus, Thyreus 259 retusa, Anthophora 256 rostratus, Thyreus 258 rowlandi, Habropoda 281 salviae, Amegilla 258 Postscript. — Since these notes were written and printed off (see p. 309), I received from Dr. I. H. H. Yarrow the following information on the type of Crocisa fulvohirta Cameron, from “Cape sapiens, Amegilla 258 sauteri, Anthophora 269, 274 sauteri, Protomelissa 269, 274 semperi, Anthophora 257 separata, Melecta 256 sibiricus, Thyreus 284, 287 sierrae, Melecta 256 simulatrix, Eupavlovskia 267 sinaitica, Paracrocisa 257, 302 (key), 305 solskyi, Anthophora 257 spinosa, Acanthomelecta 319 spinosa, Melecta 317 stanfordiana, Anthophora 259 surniculus, Thyreus 259 sutepensis, Habropoda 257, 270, 281 tainanicola, Habropoda 270 tarsata, Habropoda 256, 267 ternatensis, Amegilla 258 thoracica, Melecta 256 Thyreus 255, 258, 261 (key), 266 transcaucasica, Pseudomelecta 284, 290 transitoria, Melecta 307 tricolor, Protomelissa 257, 273 (key), 278 truncatus, Thyreus 259 turneri, Habropoda 281 urbana, Anthophora 259 ursina, Anthophora 256 verticalis, Thyreus 258 violacea, Amegilla 259 vulpecula, Callomelecta 282 vulpecula, Protomelissa 257, 272 (key) Xeromelecta 258, 259, 284 Zacosmia 259 zonatula, Habropoda 256, 267 Colony”, in the British Museum (Nat. Hist.) collection: “The specimen is a 6, not a @ as Cameron says. Gaster sternite 1 is obliquely projecting and sternites 4 and 5 have projecting bristles (I presume this is what Cameron means when he says “apices of first and of the fifth and sixth abdominal segments project obliquely at the apex”). Tergite 7 is partly projecting and its apical margin is more or less truncate but a little incised in middle. The antennae, which are complete, are short and massive, segment 2 largely hidden in apex of scape and evidently invisible to Cameron! The sculpture, colour etc. agree pretty well with what Cameron says.” TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 PLAAT 1 l 2 3 4 5 6 Fig. 1. Protomelissa insidiosa (Lieft.), & paratype from W. Java; exp. alar. 19.2 mm. Fig, 2. P. vulpecula (Lieft.), $ paratype from S. Sumatra; exp. alar. 19.5 mm. — Fig. 3. Pseudomelecta fuscipennis (Mor.), & lectotype from Bairakum; body length ca. 11.0 mm, fore wing 9.0 mm. — Q allotype from Dzhamskoe; body length 9.4 mm, fore wing 8.0 mm. Fig. 4. P. fuscipennis (Mor.), 9 Fig. 5. P. diacantha (Eversm.), holotype from “Alp. Sibir”. — Fig. 6. P. chalybeia spec. nov. 9 paratype from Mongolia; exp. alar. 19.0 mm. — Figs. 1—2 after Lieftinck (1944) M. A. LIEFTINCK : Old World Melectine bees TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 7, 1972 PLAAT 2 7 9 8 10 11 12 Fig. 7. Paracrocisa guilochei (Dusmet), & holotype from Tigui, Morocco; body length 12.4 mm, fore wing 10.0 mm Fig. 8. P. guilochei (Dusmet), & from Melilla, Morocco; body length 14.5 mm, exp. alar. 23.5 mm. — Fig. 9. P. sinaitica Alfken, & from Gebel Elba, Egypt; body length 16.0 mm, exp. alar. ca. 26.0 mm Fig. 10. P. kuschakewiczi (Radoszk.), 4 from Turcme- nia (NMW); exp. alar. 22.5 mm Fig. 11. Afromelecta fulvohirta (Cam.), 9 syntype E. kobrowt from Delareg, Transvaal (NRS) - Fig. 12. A. fulvohirta (Cam.), & from Annshaw, S.A.; exp. alar. 26.5 mm M. A. LIEFTINCK : Old World Melectine bees No. 2. P. J. Brakman, 1966. — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff, 1968 — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.FI. 70.—. No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. D.FI. 25.—. No. 5. C. A. W. Jeekel, 1970. — Nomenclator generum et familiarum Diplopodorum, 412 pp. D.Fl. 100.—. No. 6. J. G. Betrem, 1971. — The African Campsomerinae (Hymenoptera, Scoliidae), 326 pp., 47 figs., 6 pls. D.Fl. 120.—. 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TIJDSCHRIFT voor ENTOMOLOGIE Redactie (Editorial Board) . . . . . . A. Diakonoff, P. J. van Helsdingen, C. A. W. Jeekel, R. de Jong, J. Krikken, M. A. Lieftinck. Address . . 2 2 2 . . . . . „Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden. The Journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. It appears in separate issues, forming an annual volume of 350—400 pages. Subscription rate: D.Fl. 62.50 per volume. MONOGRAFIEEN VAN DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING Redactie (Editorial Board) and address as for Tijdschrift voor Entomologie. The Monographs are intended for the publication of larger entomological papers on a single subject and will appear irregularly. The following Monographs have been published: Hors serie: F. T. Valck Lucassen et al., 1961. — Monographie du genre Lomaptera Gory & Percheron (Coleoptera, Cetoniidae), 299 pages, 739 figs., 2 pl., map. D.Fl. 60.—. . 1. A. J. Besseling, 1964. — De Nederlandse Watermijten (Hydrachnellae Latreille, 1802) (The Hydrachnellae of the Netherlands), 199 pp., 333 figs., D.Fl. 30.—. LIFE CYCLE OF APODEMOPUS APODEMI (FAIN, 1965) (GLYCYPHAGIDAE: SARCOPTIFORMES) by F. S. LUKOSCHUS Zoölogisch Laboratorium, Katholieke Universiteit, Nijmegen A. FAIN Institut de Medicine Tropicale Prince Leopold, Antwerpen, Belgique & F. M. DRIESSEN Nederlands Instituut voor Zuivelonderzoek, Ede SUMMARY Apodemopus apodemi is a free living nest inhabitant of Apodemus sylvaticus, A. flavi- collis and A. callipides, with a short free life cycle, correlated with duration of pregnancy of the host and a long-period hypopus stage, endoparasitic in the hair follicles of the tail. Methods for rearing and data on the biology of this species are given. Adults and devel- opmental stages are described, and the systematic position is confirmed. Histological observations could not answer the question of nutrition of the tissue hypopi. The similarity of the life cycle with that of Hypodectidae species is discussed. SYSTEMATIC POSITION OF THE GENUS A podemopus FAIN The heteromorphic deuteronymphs of Apodemopus apodemi were first found in the follicles of the hairs upon the tail of Apodemus sylvaticus. The species has been placed in the genus Lophuromyopus and described under the name L. apodemi Fain (1965 c). The hypopi of Lophuromyopus and related genera live in a niche only observed in detail in the last years (Fain, 1965a, b, c, 1966a, 1967a, b, c, 1968b, c; Fain & J ukoschus, 1968). In connection with their endoparasitic way of life these genera lack clasping organs. The species are similar to the family Hypodectidae, the hypopi of which live subcutaneous in birds (Fain & Bafort, 1966, 1967; Fain, 1967a). Later on Fain (1967d) described the monotypic subgenus Apodemopus. This differs from the typical subgenus Lophuromyopus chiefly by scapular setae inserted far ahead of the sejugal furrow and by the presence of sclerotized processes dorsally on the idiosoma. Fain (1968a) succeeded in rearing hypopi of a Lophuromyopus species to the adult stage. The adults live free in the nests of rodents, for this group one of us (A.F.) had suggested the genus Grammolichus Fain (1968a). Revision of the genus Lophuromyopus thus became necessary (Fain, 1968b). In the meantime, one of us (F.L.) succeeded in rearing hypopi of A. apodemi to tritonymps and adults. These adults correspond in general to the genus. They differ, 326 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 however, in a number of important characteristics, which justify their separation (Fain, 1968b : 162). These characters are the following: Adults have Y-shaped epimerae I, in the female the vulva is inverted Y-shaped with three distinct valves. The epigynium is lying behind the sternum. The genus differs from the genus Lophuromyopus by the following characteristics : In both sexes: cuticle verrucose and little sclerotized (thick, heavily sclerotized and furrowed in Lophuromyopus); presence of only one solenidion on genu I; presence of one seta on femur IV; external vertical setae close together and behind v 7; no aerial sacs dorsally to trochanter II; absence of solenidion omega 2. In the male: Penis very short, no genital combs on the forelegs. The shape of the genital opening in the female (inverted Y-shaped with three valves) prevents the placing of the genus within the subfamily Labidophorinae. In this subfamily the vulva is always situated in the Jongitudinal axis and has only two valves. Two setae on tibiae I and II and the formation of hypopi keep it from the assignement in the subfamily Ctenoglyphi- nae. Volgin (1964) described Lophioglyphus liciosus out off nest of rodents in Russia. The only figure (dorsal view of the male) published with the description, gives no exact data, but the species seems closely related to Apodemopus apodemi. Unfortunately we were not able to borrow paratypes of Lophioglyphus liciosus for comparison. After rearing experiments we now are presenting the entire life-cycle of a Lophuromy- opinae species, with data on a peculiar biology. METHODS OF REARING AND COURSE OF DEVELOPMENT Apodemus sylvaticus populations within the Netherlands, Belgium, Germany, Austria and Switzerland, A. flavicollis in Switzerland and Austria, and A callipides in Spain (Lukoschus, 1967) are found to be parasitized to a high degree by the hypopi. Out of 154 investigated wood mice in the environs of Nijmegen, 122 proved to be infected during the month of September (1965—1968). During winter the degree of parasitism is lower, in March and April parasites are rare in gravid mice and absent in lactating mice. In the autumn mice bear from 1 to 182 hypopi per tail, mostly 20—50 parasites are to be found. On infected parts of the tail enlarged hair follicles can be observed macroscopically, the thickening elevating the posterior border of the scales. By striking over the tails distally with a scalpel or pincers, the hypopi are squeezed from the follicles under the scales. Initially we succeeded only to rear the adults (Fain, 1968 b). Now we consider the following methods to be nearly optimal. Profitable season is September—October. The storage of the dead, trapped mice should be at 4° C for 3—4 days; than the hypopi are squeezed from the follicles and a part of the tail is amputated. Little glass dishes with dried yeast and parts of the tail are used for rearing. Rims of the dishes are smeared with caterpillar lime to make escape from the dishes visible. Storing of the dishes is at 85% RH, darkness and room temperature. Under these conditions mould appears, apparently necessary for the development of the species. Hypopi with relatively short legs (Fig. 10, 12), set free as described above, appear to be inert. Two days later those hypopi, which are situated beneath hairs and at scale 327 F. S. LukoscHus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus ENORMI 2S02NII2A JO MATA [EISYe] ‘IouaA few ‘8961 ‘SMHOSONNT 8 NIVA zwapodr sndowapody ‘7 ‘SIA EJNIIND ISOINIIIA JO MITA [EINE] “MATA JESIOP ur A I—I ISFE} pure viadtay> ‘1ayUaA afewaz 8961 ‘SMHOSONNT Ÿ NIVA twopode sndowapody "1 ‘SIA 328 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 borders in such a manner that both ventral and dorsal surface come into contact, become unable to move. Preparations of these specimens prove them to be in the process of moulting into tritonymphs. Hypopi glued together by squeezed sebaceous fat and scales, or glued to the surface lying on the back, do not start development even within 14 days, but die after shrinking. First tritonymphs appear on the 3rd day (Fig. 5, 8). Frequently in fissures of yeast, less upon the tail, tritonymphs enter moulting phase on the 5th day. First adults are present on the 7th day (Fig. 1, 2). Tritonymphs and adults are active in the dishes, but do not try to escape. On the 9th day first eggs are found on the glass bottom, in smaller quantities on yeast and only rarely on the tail pieces. Isolated couples could deposit 24 eggs within 10 days, under laboratory conditions. Larvae (Fig. 3, 6) are present on the 12th, first protonymphs (Fig. 4, 7) on the 14th and first hypopi (Fig. 11, 13) on the 17th day. Free hypopi are remarkably smaller than tissue hypopi. They have relatively long legs (compare Fig. 10 and 11) and are very active. The conditions in the rearing dishes seem to be not convenient, for most of them become glued on the lime border. For rearing larger numbers of free hypopi use of glass tubes with a cotton stop was necessary. We did not succeed in observing infestation of the host by free hypopi. Behaviour of hypopi was observed under dissecting microscope upon the tails of arrested Apodemus up to 60 minutes. Larger numbers of hypopi, placed on adult wood mice without parasites in the tails, did not make efforts to enter beneath the scales. The following days hypopi were found in the fur. Up to four weeks no hypopi were observed within the follicles of tails of test mice. Parasitized wood mice trapped in winter did not reproduce in laboratorium mice boxes, their nests had no developmental stages nor adult mites, but nests of free living wood mice with nestlings in May harbour these stages. Numbers of free hypopi placed on one day old nestlings of laboratory mice and of Mesocricetus auritus did not enter juvenile hosts. Without a drop of temperature only few hypopi became tritonymphs. Even in high humidity tissue hypopi cannot hatch out of follicles of dead unproductive mice. Mould is necessary for nutrition of tritonymphs. Rearing tests without part of tails and the mould appearing in these conditions, do not produce adults. For egg production only yeast is the suitable food. DESCRIPTION OF STAGES Female. — Length of idiosoma of 10 specimens measured average 383 u (336— 423), width 240 u (224—266). Venter (Fig. 1). Cuticle soft, verrucose. Empimera I Y-shaped, coxal field III almost closed. Vulva between coxae III and IV with two stronger sclerotized valves (78 y) and a soft median valve. Two pairs of small genital suckers. Epigynium (eg) and genital apodemes (op) small, but strongly sclerotized. Anal opening ventrally at one-fourth from caudal end. Bursa copulatrix ventral subterminal. Gnathosoma with two-segmented palps and large strongly dentated chelicerae. Palpal coxae with two pairs of setae (lateral one pectinate). Palptibia with two, tarsus with one setae and three spines. Chelicerae as figured. Ventral chaetotaxy. Present are cx I 27 u, cx III 12 u, setiform genital setae g 4 6 u, 3 F. S. LukoscHus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus IAS SONIA JO MATA [UP] Ia ‘ydur{uoyrn zo ‘ç pue ‘ydurduojoid % ‘varey zo 199USA ‘€ ‘8961 ‘SNHOSONNT 29 NIVA suapody sudo lana] Moors —— gs EN i At, / 5 ; Ed | eo) IW) // | (7, CD4 4 cbs 4 em wapody *S—¢ ‘Sri 330 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 Fig. 6—9. Apodemopus apodemi FAIN & LUKOSCHUS, 1968. 6, dorsum of larva; 7, protonymph; 8, tritonymph and 9, female F. S. LukoscHus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus 331 &§ m7 pu, g p 22 n, anals 7 and 2, and finely pectinated setae a 3, a 4 20 ua 5 28 u, lateral 5 43 y, and subhumeral (sb) 44 y. Legs with five free segments and long ambulacrum with very little claw. Setae on tarsi all setiform, on trochanteres-tibiae short pectinate. Chaetotaxy: Tarsi 10—10—8—8, tibiae 2—2—1—1, genua 2—2—1—0, femora 1—1—0—1, trochanteres 1—1—1—0. Solenidiotaxy : Tarsi 2—1—0—0, tibiae 1—1-—1—1, genua 1—1—0—O. On tarsus I famulus is present, omega 2 is absent. Dorsum (Fig. 9). Cuticle soft, verrucose. All dorsal setae thick and finally pectinated. Present are v 7 in front of v e, sc à behind level of sc e, d 1—d 5, 11-14, h and multiple branched supracoxal setae. Measurements in table I. Oil gland between laterals 2 and 3. Male (Fig. 2). — Length of idiosoma in 10 specimens measured 284 x (266—314), width 176 u (157—193). Body shape more slender than in female. Cuticle soft, ver- rucose, papillae smaller than in female. Epimera I Y-shaped, coxal field III closed. Genital opening between coxal regions IV with two triangular genital apodemes. Penis (Pe) very short, two pairs of little genital suckers. Chaetotaxy of venter and dorsum like in female, only anal setae 1 are lacking. Measurements in table I. Chaetotaxy and solenidiotaxy of legs like in female, with the exception of tarsi, these only 8—8—6—6. Tritonymph. — Length of idiosoma in 10 specimens measured 353 u (271— 448), width 225 u (176—277). There is no morphological difference in tritonymphs, which include a developing male or female. Large range of variation in length is caused by mixture of male- and female-forming tritonymphs. Venter (Fig. 5). Epimera I free. Genital opening between coxae IV with two oblong oval genital valves and two pairs of disk-shaped. genital suckers. Chaetotaxy like in female, but anal setae 7 are lacking. Dorsum (Fig. 8). Chaetotaxy like in female, but all setae shorter and smaller, meas- urements in table I. Supracoxal setae smaller and less branched. Chaetotaxy of legs: Tarsi S—8—6—6, tibiae 2—2—1—1, genua 2—2—1—0, femora 1—1—0—1, trochanteres 1—1—1—0. Solenidiotaxy: Tarsi 2—1—0—0, tibiae 1—1—1—1, genua 1—1—0—0. Free hypopus. — Length of idiosoma in 10 specimens measured 183 y (174— 190), width 113 u (106—118). Heteromorphic nymph of the type of cuticole Rodentia parasites without sucker or claw-like organs (Fain, 1965). Cuticle smooth, shiny light brown. Venter (Fig. 11). — Epimera I fused in V-shape. Epimera III and IV form closed coxal fields III. Epimerites IV lying along genital opening. Genital opening with pairs of genital suckers beneath oblong-oval valvae. Anal opening ventral subterminal with two pint-like setae (d 5 ?). Palposoma with two small hairs each, without solenidia. Long pectinate v 7 translocated to venter, s 4 setiform, coxal setae I absent, coxals III only in form of rings. Cuticle wavy-wrinkled, enabling enlargement to tissue hypopus (Fig. 10). Dorsum (Fig. 13). Strongly sclerotized and punctured dorsal shield covers most parts TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. :8, 1972 332 sndoddy 2917 zo sojuaa ‘TT ‘sndoddy anssıy Jo 124U3A ‘OI ‘8961 ‘SNHOSONNT 8 Nivg tWopode sndouopody *11—01 ‘dig ba} en F. S. Lukoscuus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus sndoddy 2217 Jo ‘er pue ‘sndoddy anssn zo umsıop ‘ZI | 8961 ‘SNHOSONNT 9 NIVA tWepode sndowapody "EI—ZI ‘Fg GP 334 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 of dorsum. Soft parts wavy-wrinkled, enabling extension of tissue hypopus (Fig. 12). Vertical external setae pectinate, all other setae setiform to spine-like, also supracoxal setae. Two pairs of gland openings between humeral and lateral 1 and between lateral 2 and 3. There is no connection of apophysis of pygidial shield with dorsal shield. Legs I and II with ambulacrum and strong claw, legs HI and IV without ambulacrum and claw. Femora I and II with broad flattened seta within deep basis. Tibiae III and IV with hoe-shaped setae with three, sometimes four points within a deep basis. Solenidion omega 3 on tarsus I distinctly displaced more basally. Chaetotaxy of legs: Tarsi 6—6—9—4, tibiae 2—2—1—1, genua 2—2—1—0, femora 1—1—0—, trochanteres 1—1—1—0. Solenidiotaxy: Tarsi 2—1—0—0, tibiae 1—1—1—1, genua 0—0—0—0. Tissue-hypopus. — Hypopi pressed out of the follicles of tails have smooth, white cuticula, only legs are yellow-brown. Length of 10 specimens measured 322 y (282—370), width 183 x (163—214). Venter (Fig. 10) and dorsum (Fig. 12) show clearly stretching of soft, wavy parts of free hypopi. Protonymph. — Length of idiosoma in 10 specimens measured 178 u (154— 218), width 108 y (90—137). Body shape and soft, verrucose cuticula like in tritonymph. Venter (Fig. 4). Epimera I fused in V-shape with little sternum. Genital opening with two valves and only one pair of genital suckers. Only one pair of genital setae (g m) and three pairs of anals (a 2, a 4, a 5). All other hairs of venter and dorsum (Fig. 7) like in tritonymph. Measurements in table I. Remarkable are supracoxal setae, only two-pointed, v 7 relatively long. Chaetotaxy of legs: Tarsi 8—8—6—4, tibiae 2—2—1—0, genua 2—2—1—0, femora 1—1—0—, trochanteres 0—0—0—0. Solenidiotaxy: Tarsi 1—1—0—0, tibiae 1—1—1—0, genua 1—1—0—0. Larva. — Length of idiosoma in 10 specimens measured 174 u (160—196) width 107 u (90—120). Cuticula soft and verrucose. Venter (Fig. 3). Epimera I fused in Y-shape. Genital opening, genital setae and anal setae are lacking. Organs of Claparede present in shape of rings (R), not protu- berant. Dorsum (Fig. 6). Vertical external and supracoxal setae short and smooth. Dorsals 4 and laterals 4 are absent. Other dorsal setae pectinate and relatively long. Measurements in table I. Egg. — White with honeycomb pattern, 133 y long and 64 u wide. When mounting in Hoyer’s mixture, pattern disappears. DEPOSITION OF SPECIMENS Males, females and developmental stages have been deposited within: Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands. U.S. National Museum, Washington, D.C., U.S.A. Institute of Acarology, Columbus, Ohio, U.S.A. Zoology Department, University of Massachusetts, Amherst, Mass., U.S.A. F. S. LukoscHus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus 335 Muséum d'Histoire Naturelle, Paris, France. British Museum (Natural History), London, England. Institut de Médecine Tropicale Prince Léopold, Antwerpen, Belgium. Zoologisches Institut und Zoologisches Museum, Hamburg, Germany. Institute of Parasitology, Academy of Science, Prague, Czechoslovakia. Bernice P. Bishop Museum, Honolulu, Hawaii. Zoölogisch Laboratorium, Nijmegen, Netherlands. PARASITE-HOST INTERACTION To observe situation of hypopi i” sitw and to study mite-host interactions, infected tails have been sectioned and stained with haematoxylin-eosin. Parasites have entered distal parts of hair follicles (Pl. 1, Fig. 1) along the hair of the host (H), causing thickening of the epidermis (Ep) of hair follicle. Around hypopus (Hy) stratum corneum, which is normally thin within hair follicle (7 c), shows either hypertrophy (A c) or paraceratosis (p c). Also connective tissues (c #) are affected. This is similar to parasitism in Psorergates species (Lukoschus, 1967) and nasal mites (Kim & Bang, 1970). Strongly parasitized wood mice often lose epidermis of the tail, because of this degeneration of connective tissues. The parasite causes epidermal cells to degenerate (Pl. 2, Fig. 3). The nuclei become lighter of colour (1 c), only traces of them remain (g), for some time. In places of degenerated nuclei numerous dark coloured granulae (gr) appear (nucleous grit). In a later phase also the protoplasm degenerates (pr d). Another form of degeneration (Pl. 2 Fig. 2) is characterized by sickle-shaped nuclei (5 sh), the concave side always directed to the parasite. All these characteristics are similar to those developed by insect and mite bites (Allen, 1948; Tobias, 1949). Regarding destruction of tail epidermis and often its entire loss with its results, we consider Apodemopus apodemi to be low grade pathogenic. Discussion Our knowledge of the life cycle and the biology of genera and species with hetero- morphic deutonymphs (hypopi) living in the phoretic association on mammals, is restricted. Most of the genera and species are described only from hypopi (Fain, 1969). Review of literature on rearing techniques and life cycle is recently given by Lukoschus, de Cock & Fain (1971) in description of life cycle of Melesodectes auricularis Fain & Lukoschus, 1968. In Apodemopus apodemi a long period of follicle parasitism, rapid development and reproduction of free living forms, and enlargement of free hypopus to tissue hypopus are remarkable. Development from tissue hypopus to free hypopus occurs within a short time (shortest observed time is 17 days under laboratory conditions). Even if devel- opment would be distinctly longer under field conditions, there will be sufficient time for infection of nestlings, because gestation and lactation periods in wood mice are remarkably longer. Tissue hypopi seem unable to free themselves from the follicles. This is similar to what is observed in genera of the subfamily Hypodectinae, parasites of birds. Devel- opmental cycle has been observed by Fain & Bafort (1966) in Hypodectes propus 336 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 Nitsch, 1861, subdermal parasite of pigeons. In this species the large tissue hypopi are eliminated by the bird. This is suggested to be affected by hormonal conditions of the bird while breeding. Hypopi give direct development to adults, which do not feed. From the numerous eggs hatch small free hypopi (larval and protonymphal stages are only rudimentary within the egg shell). Free hypopi enter the young nestling and extend 7—10 times. In A. apodemi similar enlargement occurs (compare Fig. 10 and 11), but to a less extent. In this species feeding is not restricted to hypopial stage only, like in Hypodectes, but free living stages also need appropriate food. Yeast, a suitable nutrient for free living mites, is sufficient for reproduction and development of free hypopi, but is not sufficient for development of adults. We are not sure whether mould alone, rising from the tail parts in rearing dishes, contains all factors needed for development, or whether also debris from the host, available on the tails in the rearing tubes, are necessary. Host-born debris is present in nest under field conditions. Dependence on hostal debris has been observed in Melesodectes auricularis and Marsupialichus marsupialis. Development into the adult stage and reproduction seem to be very variable according to the genera observed. Fain (1968b, 1969b) succeeded in rearing hypopi of Lophur- omyopus and Rodentopus without food to adults, but obtained no eggs and no further development. Working with Labidophorus talpae and Orycteroxenus dispar, Fain (1969c) succeeded in obtaining tritonymphs, but was unable to rear adults from the latter. In Dermacarus sciurinus Koch, 1841, we succeeded to get a whole developmental cycle by storing tails of the host at 85% relative humidity. Feeding of hypopi without mouth parts is still an open question. Hypopi enlarge enormously within tissues, and tissues show characteristic degenerations, as observed from stinging insects and mites possessing salivary glands with extraintestinal digestion. It may be suggested that enormous dorsal glands act in such a way; however, the place of osmotical absorption is still unknown. BIBLIOGRAPHY Allen, A. C., 1948. Persistent “insect bites” (dermal cosinophilic granulomas) simulating lymphoblas- tomas, histiocytosis, and squamous cell carcinomas. Am. J. Pathol. 24: 367—388. Fain, A., 1965a. Quelques aspects de l’endoparasitisme par les acariens. Ann. Parasitol. 40: 317— 327. ——, 1965b. Un nouveau type d’hypope, parasite cuticole de Rongeurs Africains (Acari : Sarcopti- formes). Z. Parasitenk. 26: 82—90. ——., 1965c. Nouveaux genres et espèces d’Acariens Sarcoptiformes parasites. Rev. Zool. Bot. Afr. 72: 252—256. ——, 1966. Note sur les Acariens nidicoles à deutonymphe parasite tissulaire des Oiseaux (Hypodectidae : Sarcoptiformes). Rev. Zool. Bot. Afr. 74: 324—330. ——., 1967a. Les hypopes parasites des tissus cellulaires des oiseaux (Hypodectidae : Sarcoptifor- mes). Bull. Inst. r. Sci. nat. Belg. 43: 1—139. ——, 1967b. Nouveaux hypopes vivant en association phorétique sur les rongeurs et des Marsu- piaux (Acarina : Glycyphagidae). Acarologia 9: 415—434. ——, 1967c. Les Hypopes des Glycyphagidae nidicoles en Afrique au Sud du Sahara. Ann. Mus. roy. Afr. centr. Tervuren 157: 1—89. ——., 1968a. Acariens nidicoles et détriticoles en Afrique au Sud du Sahara. III. Espèces et genres nouveaux dans les sous-familles Labidophorinae et Grammolichinae. Acarologia 10: 86—110. ——, 1968b. Acariens nidicoles et détriticoles on Afrique au Sud du Sahara. IV. Découverte du cycle évolutif d'un Acarien du genre Lophuromyopus FAIN, 1965 (Glycyphagidae : Sarcopti- formes). Rev. Zool. Bot. Afr. 78: 161—174. Fain, A., Fain, A. Fain, A. Fain, A. F. S. LukoscHus, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus 337 1968c. Un hypose de la famille Hypoderidae Murray, 1877 vivant sous la peau d'un rongeur (Hypoderidae : Sarcoptiformes). Acarologia 10: 111—115. 1969a. Les deutonymphes hypopiales vivant en association phorétique sur les mammiferes (Acarina : Sarcoptiformes). Bull. Inst. r. Sci. nat. Belg. 45: 1—262. 1969b. Acariens nidicoles et détriticoles en Afrique au Sud du Sahara. V. Description du cycle évolutif de deux espèces du genre Rodentopus, Fain. Acarologia 11: 304—316. 1969c. Morphologie et cycle évolutif des Glycyphagidae commenseaux de la Taupe, Talpa europea. Acaologia 11: 750—795. & J. Bafort, 1966. Les hypopes parasitant les tissus cellulaires sont les deutonymphes d'un acarien libre et pas celles d’un acarien plumicole. Rev. Zool. Bot. Afr. 74: 315—316. & J. Bafort, 1967. Cycle évolutif et morphologie de Hypodectes (Hypodectoides) propus (Nitzsch) acarien nidicole 4 deutonymphe parasite tissulaire des pigeons. Bull. Acad. Rov. Sci. Belgique, 5. Sér. 53: 501—533. & F. Lukoschus, 1968. Une nouvelle deutonymphe hétéromorphe (hypope) parasite du blaireau (Meles meles) en Hollande (Acarina : Sarcoptiformes). Rev. Zool. Bot. Afr. 78: 175—182. Kim, C. S. & B. G. Bang, 1970. Nasal mites parasitic in nasal and upper skull tissues in the baboon (Papio sp.). Science 169 : 372—373. Lukoschus, F., 1967. Krätzmilben an spanischen Kleinsäugern. Rev. Iber. Parasitol. 27: 203-228. Lukoschus, F. S., A. W. A. M. de Cock & A. Fain, 1971. Life cycle of Melesodectes suricularis Fain & Lukoschus, 1968 (Glycyphagidae : Sarcoptiformes). Tijdschr. Ent. (in press). Tobias, N., 1949. Tickbite granulema. J. investigat. Dermatol. 12: 255—259. Volgin, V., 1964. New genus and new species of mite of the subfamily Ctenoglyphinae (Glycyphagi- dae) from the nests of Rodents. Acad. Nauk. S.S.S.R. Parasitogieski sbernik 22: 100—110. 338 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 Table I. Length of setae. Heasurements in u. ui ' Li LI 1 ! 1. !trito- ! ! proto- male female nymph hypope nymph re AA eee ee lee ed coxal I 18 Dili 15 - 11 coxal III 18 12 8 0 genital anterior 11 6 6 6 genital median 10 Fi F4 - genital posterior 15 22 6 - anales 1 - 7 - - anales 2 8 9 5 = anales 3 8 A 6 = anales 4 35 28 13 = 10 anales 5 25 28 18 - vertical internal 49 50 41 24 34 vertical external 80 94 28 19 12 scapular internal 74 106 42 9 Bal scapular external 12 81 52 9 30 supracoxal 18 18 13 5 humeral 59 69 45 5 18 subhuneral 41 44 23 6 17 dorsal 1 66 delig, 50 8 34 dorsal 2 67 1 107 52 6 29 dorsal 3 74 ES 54 6 29 dorsal 4 67 120 46 6 32 dorsal 5 51 65 39 ©) 22 lateral 1 62 74 41 7 30 lateral 2 64 93 41 6 24 lateral 3 67 83 43 6 24 lateral 4 60 65 41 5 18 lateral 5 41 43 36 6 14 solenidion phi I 170 106 83 33 60 solenidion phi II 98 70 54 21 33 solenidion phi III 21 21 18 5 solenidion phi IV 5 5 3 5 solenidion omega; I 9 10 9 6 solenidion omegaz E 20 31 20 1112 length tarsus I 49 46 33 25 23 length tersus II 39 Ab 31 24 20 length tarsus III 44 59 35 48 27 length tarsus IV 54 78 36 15 31 F. S. LUKOSCHUS, A. FAIN & F. M. DRIESSEN : Life cycle of Apodemopus 339 EXPLANATION OF PLATES Plates 1 and 2. Longitudinal sections of Apodemus sylvaticus tails parasitized by Apo- demopus apodemi. Bouin-fixation, 7 p-sections, haematoxyline-eosine-staining. | Plate 1 Fig. 1. Parasitised hair follicle. Plate 2 Fig. 2 and 3. Parasitised hair follicle, details of epidermis. REGISTER VAN DEEL 115 * Een sterretje duidt een naam aan nieuw voor de wetenschap. * An asterisk denotes a name new to science. Synoniemen zijn cursief gedrukt. Synonyms are printed in italics. Uit dit register zijn weggelaten de namen voorkomende in artikelen welke reeds werden voorzien van eigen registers, nl. die van sommige Lepidoptera (pp. 1—121, index p. 121), Hymenoptera Sphecidae (pp. 153—203, index p. 203), en Hymenoptera Apoidea (pp. 253—324, index p. 323), in dit deel. ACARI apodemi 325 et seq. Apodemopus 325 et seq. Dermacarus sciurinus 336 Grammolichus 325 Hypodectes propus 335 Labidophorus talpae 336 Lophioglyphus liciosus 326 Lophuromyopus 325, 336 LEPIDOPTERA Allohermenias 246, 247 Argyroploce 243 azukivora 243 Cryptophlebia 241 *elpisma 242 elutana 243, 245 Eucelis 250 Eucosma 243, 244 Marsupialichus marsupialis 336 *felix 243 Melesodectes auricularis 335 Orycteroxenus dispar 336 Psorergates 335 Rodentopus 336 ARANEAE Pardosa 205 et seq. prativaga 205 et seq. pullata 205 et seq. sphagnicola 205, 206 AVES Lyrurus tetrix 150 Philomachus pugnax 150 DIPTERA chrysorrhoea 137 Physocephala 137 HYMENOPTERA Andrena 123 Dasypoda 123, 128 Halictus 123 Hedychrum 134, 138 hirtipes 128 intermedium 134, 138 Megachile 123 Philanthus 123 et seq., [141 et seq. Laspeyresia 250 Lathronympha 243 Matsumuraeses 241 et seq. melanaula 243, 248 metacritica 241, 246 monstruosana 242, 243 ochreocervina 241, 250 phaseoli 241, 243, 248, 251 Semasia 241, 243 *tetramorpha 247, 251 Thiodia 243 trophiodes 243, 245 *xantholoba 245 MAMMALIA Apodemus callipides, flavicol- lis and sylvaticus 325, 326 Mesocricetus auritus 328 ODONATA albinensis 224 Agriogomphus 218, 220 anduzei 227 Aphylla 222—226 Archaeogomphus 218 argentina 230 batesi 236 *boliviana 222 calippus 237, 238 conchinus 221 Cyanogomphus 220, 221 dentata 224 Desmogomphus 221 340 distinguenda 226 Epigomphus 221 ericae 220 eugeniae 233 gladiata 228 jessei 218, 220 klugi 238 lieftincki 237 llama 221 modesta 227 molossus 225 nanus 218 Negomphoides 236 pallida 236 paucinervis 221 Phyllocycla 223, 227—236 *propinqua 235 spectabilis 237 surinamensis 238 sylvicola 220 theodorina 226 tigrivensis 221 titschacki 227 tumens 220 uncatus 221 undulatus 236 * vesta 227 viridipleuris 233, 235 Zonophora 236—238 PLANTAE Astragalus membranaceus 251 Cajanus 248 Crotalaria 242 Erica cinerea 123 Glycine Max 245 Hydrocotyle vulgaris 206, [207, 210 Lysimachia vulgaris 206, [207, 210 Phaseolus 248 Pinus pinaster 123, 141 Polytrichum 206 Tephrosia vogelii 245 Ulex europaeus 123, namis 123 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 PLAAT 1 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 115, AFL. 8, 1972 PLAAT 2 F. S. Lukoscuus et al.: Life cycle of Apodemopus No. 2. P. J. Brakman, 1966, — Catalogus van Coleoptera uit Nederland en het omliggende gebied (List of the Coleoptera of the Netherlands and adjacent region), 219 pp., map. D.Fl. 45.—. No. 3. G. A. Graaf Bentinck en A. Diakonoff, 1968 — De Nederlandse Bladrollers (Tortricidae), (The Netherlands Leaf-Rollers (Tortricidae) ). 350 pp. en 98 pl. D.Fl. 70.—. No. 4. F. Willemse, 1968. — Preliminary revision of the genera Stenocatantops Dirsh & Uvarov and Xenocatantops Dirsh & Uvarov (Orthoptera, Acridiidae Catantopinae), 77 pp., 1 map, 6 pl. D.Fl. 25.—. No. 5. C. A. W. Jeekel, 1970. — Nomenclator generum et familiarum Diplopodorum, 412 pp. D.F]. 100.—. No. 6. J. G. Betrem, 1971. — The African Campsomerinae (Hymenoptera, Scoliidae), 326 pp., 47 figs, 6 pls. D.Fl. 120.—. 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