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V

A REVISION OF THE GENUS CALLIP TAMUS SERVILLE

(ORTHOPTERA : ACRIDIDAE)

BY

N. D. JAGO Univenity of Ghana. Accra

Pp. 287-350; 26 Text-jigtcres

BULLETIN OF THE BRISISH MUSEUM (NATURAL HISSORY)

ENSOMOLOGY Vol. 1 3 No. 9 LONDON: 1963

T H E B U L L E T I S O F T H E B R I T I S H B I U S E U M ( N A T U R A L H I S T O R Y ) , institrited in 1949, is isszted i i t jizte series, correspondillg to the Departments of the Alirsezm, niid m z Historicnl series.

Parts Lcill appear nt irregzrlnr iiitervals as they become retidy. Vo l imes vi l1 coiztaiit aboitt three OY f o w htcndred pnges, nitd will not itecessnrily be cmnpleted withilz o.ne c n l e n h r year.

Tltis paper is Vol . 13, Xo. g of the Entomological series. T h e nbbreviated titles ojperiodicals cited follou those of t k a WorId List of Scientijic Periodicals.

0 Trrictees o[ the British Miiseuni 196.3

P R I N T E D B Y O R D E R O F T H E T R U S T E E S OF T H E B R I T I S H i M U S E U M

~ s s l l d j 1 .\I(Z)' 196.3 Price Ti;w+v-tz;.o Sltillings

A REVISION OF THE GENUS CALL IP TAM US SERVILLE

(ORTHOPTERA : ACRIDIDAE) By N. D. JAGO

CONTENTS Page

INTRODUCTION . . 289 MATERIAL . . 292 TREATMENT 294 ACKXOWLEDGEMENTC . * 294 KEY TO THE GENERA OF THE SUBFAMILY CALLIPTAMINAE . . 295 CALLIPTAMUS Serville, 1831 . 29s

S Y N O P S I S

The trans-Palaearctic genus Callipíamus Serville is revised, thirteen species now being included in the genus. The genus consists of two main elements, a northern temperate group of four species and a southern ternperate group of nine. The genus Metromerus Uvarov is synonymized with Calliptamus. A provisional key to genera in the sub-family Calliptaminae has been drawn up, together with keys to species and subspecies in the genus Cailiptamus. Observations are given on polymorphism in the genus, geographical vanation, and posible correlation of variation with climatic factors. Two species are newly described.

INTRODUCTIOS

Calliptamzts Serville, 1831, with Gryllzcs Loczcsta itnlicus L., 1758 as type, originally contained two additional species now known as ArcyPtera ficsca (Pallas, 1773) and V d e r m aeneo-oculata (De Geer, 1773). In 1838 Serville added C. ictericzis and C. margilzellzcs to the genus, though omitting mention of Acridium barbarzcm Costa, 1836. His descriptions were based on Spanish material, and it ic clear that he confused C . italiczcs (L.) (which does not occur south of the Pyrenees) with C. wattenwylianus (Pantel), “ ictericzcs ” being erected for C. bnrbnrics (Costa) material coliected near bladnd. Early in 1838, the genus under its emended form of Caloptelzus (created by Burmeister), contained the type of this genus and four other species now known as

phjmits haemntopiis (L., 17 j8), and Rnchitopis melnnoplzis (Burmeister, 1838). Later in 1838, Burmeister introduced Caloptews sicdits, clearly synonymous tvith C. bnrbnrzis (Costa). Wder geographical data were @ven by Fischer (18j3), his genus including Cnlopferttis sictclus, barbarus, ifaliczcs, and ictericzcs. C. barbarus he claimed intergraded nith italiczis, while the latter was described as possessing a variety 9nnrgiizellrcs. I t may be noted that Senille suggested varietal status for iiiarginellzts when he described it in 1838, 1. Bolivar (1876) later endorsing this view.

\

4 t Sfelnnoplzrs fenticr rubriint (De Geer, 1773), Melnnoplus bz’vittatzcs (Say, 183 j), EUYY-

24 ESTOM. 13, 9

290 ?;. D. J n G O

Walker (1870) described CaZupteizzrs discoidalis as new, and placed italiczcs (L.) and icfericzis Serville in Caloptenais : he probably confused his species with barbnrzis (Costa) and ~ ~ a t f e ~ t ~ y l i a s z i i s (Pantel) as Serville had done earlier, since his “ ifaliczis ” occurred in Syria and Israel, his “ ictericzis ” in Sorth Africa. The emended name Calliptelzzis \vas given to the genus by Sta1 (1873). The type of the genus and five other species were included, the modern Splzodmnerzrs sernpis (Serville, 1S38), Plegmatopteriis iriszis (Serville, 1838), Eziryphymzrs hnematopzis (L., I758), Calliptamiczu semirosezs (Senille, 1838), and Arceziryí&hynzzts eryflzropzis (Thunberg, 181 j). 1. Boli- var (1876), writing on the Orthoptera of Spain and Portugal, mentioned C. italiczts (L) and C. ictericris (Serville), the fonner probably being C. mzffenwylialzzrs (Pantel) and the latter C. bavbarzis (Costa). Since the name C. ictericzis was also applied later to material of C. wattenwylianzu (Pantel), e.g. by Brunner (1882), it is difficult to interpret Bolivar’s species. Brunner, in 1882, started a trend by synonymizing ali previously recopized species and subspecies under C. italiczis (L.), his Caloptenzrs brzinlzeri Sta1 later being made type for the genus Pnracaloptelztcs 1. Bolivar. As late as 1898 1. Bolivar stili agreed with Brunner in giving C. wattevzwylialzzis vanetal status

Kirby (1890) clarified the correct generic synonymy, and fured C. italicus (L.) as logotype. In 1902, Jacobsen & Bianchi put the genera Sphodromerus, Calliptamzcs, and Paracaloptelztcs into a group, the Calliptamini. Caloptelzzis coelesyrielzsis (Giglio- Tos, 1893) \vas added to Calliptamzcs by them, though siczrlzrs Burmeister and wattelzwylia.nzs (Pantel) retained their varietal status. To the varieties of Caloptemis barbarzis, Vosseler added var. deserticola in 1902.

Kirby’s catalogue of 1910 synonymized most of the post-Linnaean species under Callifitamzis italiczis (L.), six other species being included in the genus CalliPiamus : discoidalis Walker, marmarotzis Fischer-IValdheim, cephalofes Fischer-Waldheim, vzrlca.nizts Krauss, tarsizis Fischer-Waldheim, and sczitz’fer M’alker. Of the first four, zuclcanizis was given its correct name of C. plebeizis (Walker) in 192 j, and the others synonymized under C. barbnrzis (Costa). C. tarsizrs and C. sczitifer were eventually removed from the genus.

Recognition of C. bimaculatzu lirauss, 1902 (now in the genus Cdoptefiopsis)l C. abbreviatzis Ikonnikov, 1913, and C. okbaensis Kheil, 1915, started the last phase in taxonomic development, in which the numbers of species and subspecies in the genus greatly increased. Between 1928 and 1940, however, 8 new species and 4 subspecies appeared. In 19 ji, Ramme descnbed C. italiczis ilzszrlaris., C. temicercis syriaczrs, C. telzuicercis azwnlztipes, C. barbnrzis pallidipes, and C. palaestinensis erythrocnemis, Maian naming C. t em i - cercis iraciis and Mishchenko C. barbarzis italztis. Also in 1951 Nishchenko described two new subspecies of Aletromerus coelesyriensis (Giglio-Tos) : ilzfricntris and hissaricus. In rg j2, C. nfg?mtzcs Ramme and C. bnrbarzispnllidipes f . salilza hlaran were described.

The basis of the following revision is a comparative study of the male phallic complex. This provides good species and subspecies characters. The components of the phaliic complex can be seen in Text-figures I and 2, orientation irc sifci within the insect being demonstrated in Text-figure 2: J, p. 293.

The nomenclature of the phallic comples is largely that followed by Dirsh (1956)

ody.

Uvarov (1922) still recognized only two species.

. v

29 1 A R E V I S I O S OF T H E G E S U S C A L L I P T A M U S S E R V I L L E

(for key to abbreviations see Test-figure 1). Xot ali the phaiiic cornponentc are of taxonomic value. The epiphallus, while poscessing characterc of genenc importance, can be very variable (see Test-figure I j: Jr, o, Q. p. 31 j). So too are the cingular rami (Text-figure Ig) and anterior espansionc of the penis valves (Test-figure Ih). BIost ctability is found in penis and cingular valve morphology, to which aii other characters (phaíiic or esternal) can be related. Within the genuc aedeagal types faii into two groupc :

f

b

k FIG. I . Esploded generalizeci diagram of male genitalia in genus C a l l i 9 t u n ~ s Serville.

a. arch of cingulum and median dorsal cingular valve ; b. zygoma, a transverse dorsal part of cingulum to u-hich arch of cingulum and cingular valve are joined by membrane ; C. dorsal part of ectophallic membrane ; d. epiphallus ; e. oval sclerites of epiphnllus ; f. cingular apodeme ; g. ramus of cingulum ; h. interna1 anterior espansions of penis valves, for muscle attachment ; i. ejaculatory ducts ; j and k. endophallic sacs ; 1. lateral appendices of penis valves ; m. apes of penis valve. Tlie above letter key has been used throughout the paper for al1 phallic diagrams.

a92 K. D. J A C O

(i) Those with elongate blade-like penis valves, whose membranous lateral appendices curl upwards over the tongue-like cingular valve, e.g. C. ifnlicris (L.) (Test-figure 10: c and D. p. 307). The group contains the northern temperate element of the genus, best adapted to cooler, wetter, boreal conditions. Of the four species in the group, C. abbveviatzts Ikonnikov has probably evolved from C. ifaliczts (L.) under isolation in the Far East and central Asia, while C. tzirarzicrrs Tarbinsky has evolved from C. wattenuyliams (Pantel) when the two species were separated by the unfavourable semi-desert conditions which now prevail in the Bliddle East (Test-figures 6 and 7,

(ii) Those in which the penis valves are blunter, often rugose and deeply pigmented, the lateral appendices of the penis valves merely abutting on the cingular valve laterally, the appendices being thickened and frequently auricular in shape. The cingular ralve is broad and apically thickened, the posterior edge usuaiiy being straight in outline, e.g. C. bnrbnvtis (Costa) (Text-figure z j: c and F. p. 339). The group forms the southern temperate element of the genus, best adapted to warmer, drier, semi-desert or dry hlediterranean conditions. Of the nine species in the group, C. bnrbarus (Costa) has widest distnbution, and has given rice to two island species (C. madeirne Uvarov and C. plebeizis (Walker)), tvhile three further species may have arisen from C. barbarzts as a result of the isolation of elements during the last glacial espansion in Europe (C. sttbnlpirzzrs sp. n., C. cyremzicczu sp. n., and C. siciliae Ramme). C. coelesyriensis (Giglio-Tos), C. temiicercis (Tarbinsky), and C. balziclzn Uvarov complete the group, C. bnlzrcha probably having evolved in the isolation of the Western Himalayas from elements of C. terzziicercis. All subspecies in the group seem to be centred on moun- tainous areas where a rigorous climate and isolation in a rugged topography have necessitated marked evolutionary change. Thus C. barbnrzis $Jn¿aestimwsis (Ramme) is centred on the mountain backbone of Lebanon and Israel, C. bulucha braclzypterus (Dirsh) on the higher valleys of the Hindu Kush, and C. coelesyrierzsis hissnrzcus Siishchenko on the Tien Shan and mountains of northem Afghanistan.

Phaiiic characters have been used for this group by Tarbinsky (1g30), Silvestri (1g34), Grassé & Hollande (1g4j), Chopard (1943), and Ramme (rgjI). Unfor- tunately they deal in each case escept the last with only a s m d range of the distribution of each species.

PP. 301,302).

31 ATE R 1 AL

In addition to the collection of material at the British Museum (Natural History), types and coliections were made available by the following institutions through the courtesy of the specialists listed in the acknowledgments section : PrivodnajaEki JIusej Srpske Zemlje, Beograd ; Zoological Institute, Academy of Sciences of U.S.S.R., Leningrad ; Museo Civico di Storia Katurale, Genova ; RIuséum Sational d’Histoire Naturelle, Pans ; Laboratoire Evolution, Université de Pans : Istituto di Zoologia, UniversitA di Kapoli ; Osservatorio Fitopataiogico, Genova ; Department of Entomology, Sational RIuseum, Praha ; Instituto Español de Entomologia, Madrid ; The Hebrew University, Jerusalem ; Locust Control, Ministry of Agri- culture, Damascus ; Zoologisches Museum of the Humbolt-Universitat, Berlin ; The Anti-Locust Research Centre, London.

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A R E V I S I O S O F T H E G E K U S C A L L Z P T . 4 111 C S S E R V I L L E 293

, ,

E G

F H

FIG. 2. A. hhie genitalia of c. xwfíeizrryflinmrs (Fantei) i;¿ sitik with ectophailic pocket drawm back revcaiing apices of penis valres. ((1. supra-anai piatz ; p. triangular parameres eacli ui th a sensory style apicalip) ; B.-H. Iiirier stirface of posterior femora in variouc specitticris of C. bavbnvlrs (Costa) and C. itniicrrs (L.) : E-1). C. barbaruspalae- stzneizsis liarnnic Israel, Jeriisalem ; F-G. C. bnubcruirs bnrúnrlrs (Costa) from Banvuls- sur-hIer, easterri Pyreneec, siiouiri: range of variatiori for comparison with *pica1 specimen of C. ,¿fcrlicirs (L.) frorii the same area (E) : J . h longitudinal section of male Callipfatnirs type male genitalia i t t situ (compare with Figure 1 ) . Cymbois as for fig. I.

294 h-. D. J A C O

T R E A TM E S T

Under each species type data and despository foliow the name, while the diagnosis which follows the synonymy deals jn tum 111th the morphology of the male phallic complex, cercus and tegrnina, general colouring, posterior femorai and tibiai colour, hind wing morphology and colour. Females are discussed separately and their morphology compared with that of the males. The wing vein nomenclature used is that of Ragge (1955). XIeasurements are given in millimetres throughout, the number of individuals measured, range and mean, being listed for males and females. The following measurements are used in this paper.

This applies (i) Total length-distance from frons to apices of folded tegmina. even in brachypterous forms.

(ii) Head width-horizontal distance across outside of compound eyes. (ui> Femur length-distance from proximal end of posterior femur to tip of outer

apical lobe at joint with tibia. The term “ knee of posterior femur ” refers to expanded distai portion.

(iv) Tegminal length-distance from intersection of costal border of tegmen with posterior edge of pronotum to apex of tegmen, wings and tegmina being folded.

Sections on distribution and material examined are followed by a discusion section, keys to subspecies being inserted where appropriate. All material examined is represented by series or specimens in the collections

of the Bntish Museum (Natural History), uniess otherwise stated. Aü types mentioned in the text were examined by the author, or, if the holotype was not available, representative valid paratype material was used.

Keys to species proved especialiy difficult for females, where colour characters were of necessity one of the main criteria for separation. Two keys are therefore included, and that to males should be used for diagnosis wherever posible. Isolated female material may be tentatively identified from the key (p. gro), which is geographically sub-divided.

Full details of material under “material esamined” sections are available in the thesis held in the library of the University of London, Senate House, London, W.C.I.

Wherever the abbreviation C. is used it stands for CaZZiPtamzcs. Other generic emendations or synonyms beginning wlth C are written in f d .

Two references may sometimes be given under one name in the synonymy sections, e.g. see p. 333. The second, after a semi-colon, refers to an important subsequent refer- ente to the earlier name.

c

C ACKSOWLEDGEJIEXTS

This revision has been made posible under a recearch grant @ven by the Anti- Locust Research Centre, London. Special thanks must go to Sir Bons Uvarov, former director of the Centre, Dr. V. Jí. Dirsh, and the Centre’s library staff, for their help in prepanng this paper, both in supplying records and in providing advice. Access to the coliections at the Britich Museum (Natural History) was facilitated through the then Keeper of Entomology, Dr. \Y. E. China, and Dr. D. R. Ragge

A R E V I S I O S OF THE G E S U S C A L L Z P T A M U S S E R V I L L E 295

in charge of the Orthoptera section. The following ccientists supplied invaluable material, either as gifts or on loan (e.g. types) :

Dr. 2. R. AdamoviiC (Beograd), Prof. Dr. G. >'a. Be;-Bienko (Leningrad), Dr. F. Capra (Genova), Prof. L. Chopard (Pans), Prof. P. P. Grassé (Paris), Dr. 31. La Greca (Napoli), Prof. G. Jannone (Genova), Dr. J. Maian (Praha), Dr. E. Morales Agacho (ItIadnd), Dr. L. L. Alishchenko (Leningrad), Dr. 31. P. Pener (Jerusalem), Prof. E. Séguy (Paris), hfr. R. Skaf (Damascus). Dr. J. Klapperich supplied very valuable Afghanistan series, while Dr. La Greca aliowed me to have a preview of his most recent paper on the ecology of Calliptatnzcs in the Italian peninsula.

Finally Prof. O. W. Richards of Imperial College, London, and Prof. E. E. Edwards of the Cniversity College of Ghana, must be thanked for consenting to supervise this research.

I ~ E Y TO THE GENERA OF THE SUBFAUILY CALLIPTAJIISAE

The eleven genera in the Calliptaminae are all related on epiphallic characters (Dirsh, 1956), and possess cerci modified in some degree as clasping organs. The genera can only be defined using a number of characters. Uvarov (igjo) recognized the provisional nature of his key, and explained that the characters used were probably too arbitras. to be retained. Examination of Cnloptenopsis 1. Bolivar indicated that it is probably a compound of five genenc units, at least one of which is inseparable from Acorypha Krausc.

A bnef review of genenc characters follows : (i) Cerczrs-weak and tapered, e.g. Palaciosa C. Bol. and Indomevzts Uv. (Test-fig. 3 : P and

Q) , o í powerful and tri- or bi-lobed, e.g. Calliptamus Serv. (Text-fig. 3 : O) ; usually bearing an inwardly directed hook apically (Text-fig. 3 : N and O). In Palaciosa weak with no such hook. In Calopttwaopsis 1. Bol., Boihrocaracvis Uv., Paracaloptenzis 1. Bol., and Acorypha Kr., shorter than that in Calliptamus Serv., biiaterally greatly flattened, and broadly lobed apically. Ghanaian material of Stobbea riggenbachi Rme. shows a distinct trend towards formation of an apically tri-lobed tq-pe. The tendency is thus not confined only to Cnlliptamzrs Serv.

(ii) Posterior tibia1 spzrrs-in Cnlliptamrcs Serv. (Test-fig. 20 : E. p. 3 ~ 4 ) ~ outermost and longer of inner pair is unmodified. Calopenopsis 1. Bol. and Acoryphn Kr. show a tendency towards elongation of this spur, its tip finally becoming pre-apically placed behind a hirsute lobe (Test-fig. 3 : U, and Text-fig. 20 : F and G. p. 324). One Acovypha species in Ghana has both inner spurs modified. Botlwocavacvis Uv., and to a lesser estent Itzdomerrrs Dirsh, show modified spurs. The character may not indicate phyletic relationship.

(iU) Tegmina-in Calliptamzu Serv. all intermediates occur between a fully alate state and an extreme brachypterous condition (where tegmina become transformed into dorso-laterally situated scales. In contrast al1 Paracaloptenus 1. Bol., Palaciosa C. Bol., Indomerrrs Dirsh, a d Peripolrrs Mart. species are characterized by scale-like tegmina.

(iv) Pronotzrm-many Caloptetropsis species resemble Callipplamus Serv. in having a weakiy developed median carina, and slight anterior convergence in lateral carinae (Test-fig. 19 : EJ. p. 322). Typical Acovypha Iír. species have strongly convergent lateral carinae and an acuteiy pointed median posterior border, many " Cnloptenopsis " species iinfortunately showing strong trends towards this Qpe. Boszmtia Rme. and Bvachyxetzia Kirby have lateral carmae almost obliterated, and a broad pronotum with moderate to heavy surface rugosiq. Pevipolrrs Mart. and Paracaloptenus 1. Bol. have strongly tectiform pronota with smooth flat inter-carinal areas, distinct carinae, and a facies similar to that of the unrelated West African genus JIaznen St%i. Much apparent siniilarity may be due to convergent evolution.

24s EXTOJI. 13, 9

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K . D. JACO

6-9 A

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F

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B fll C D E

G H J

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R S T U

A RET’ISION OF T H E G E S U S C A L L I P T A M U S S E R V I L L E 297

(v) :Vale phaZlic complex-if this complex organ can be taken as being a more reliable indicator of phyletic affinity, then Brachyxenia Kirby and Peripolus hfart. (Text.fig. 3 : C and D) show strong resemblances, as does Indomerus Dirsh, with C. coelesyriensis hissaricus (Mistsh.) (Text-fig. 3 : A, and 14: B. p. 313). Cnloptenopsis 1. Bol. seems to be a compound genus. some groups of species resembling Acorypha type in phallic structure (see Text-fig. 3 : L, and Text-fig. 3 : E and G-J). Palaciosa C. Bol. is unique (Text-fig. 3 : B). I t is of course difficult as yet to homo- logize parts in different penis valve systems.

The following key is based mainly on male facies. 1. Cingular valve with edges curled inwards forming a partly closed tube (Text-fig. 3 : C

-. Cingular valve flattish or triangular in cross section, not forming a tubular structure

2 . Pronotum smooth and strongly tectiform with 3 distinct carinae. Hind femora slender. Tegmina forming dorso-laterally situated scales. Cerci apically bilobed-lower lobe longer than upper . . . PERIPOLUS Martinez

-. Pronotum rugosely sculptured with almost obliterated lateral carinae. Hind femora toothed above, hairy below; very broad. Cerci bilobed : lobes roughly equal oí upper longer than lower . . BRACHYXENIA Kirby

3, Lateral accessory processes of penis valves absent. Cingular and penis vaives elongate, blade-like, and of same length. Cercus very weak ; apically bilobed (Text-fig. 3 : P. p. 296) . . . . PALACIOSA C. Bolivar

-. Lateral accessory processes of penis valves well developed. Cercus moderately to

4. Vertex of head with two depressions, an anterior oval depression and a narrow inter- ocular depression. Frons espanded anteriorly, when head viewed from a lateral ‘aspect. Cercus bilobed apically with small ventral lobe similar to Caloptenopsis. Outermost of inner pair of posterior tibial spurs with pre-apical tooth (as in Text-fig. 2 0 : F. p. 324) . . B O THR O C A R A C R IS Uvarov

-. Vertex of head with single shallow ovate depression (especially in 9) or a single inter- ocular groove. Frons not unusually expanded anteriorly. Cercus bi- or tnlobed

5 . Cercus rugosely pitted on outer surface, rather weak and markedly tapered apically. Apex bilobed (Text-fig. 3 : Q. p. 296). Penis valves well developed (Text-fig. 3 : A. p. 296) but unlike most of genus Calliptamus, flattened into a plane parallel with that of cingular valve. Unlike C. coelesyriensis hissaricris (31 ishch.) in having penis valves chorter than lateral accessory processes. Cingular valve narrower than distance between outer edges of penis valves . INDOMERUS Dirsh

and D. p. 296) . 2

(Text-fig. 3 : A-B, E-M, and 10 : E-G. pp. 296 and 307) . 3

strongly developed. Bi- or trilobed apically . 4

apically. Posterior tibial spurs modified or normal . 5

I

t Frc. 3. A-M. Penis valves of representative species from genera in sub-famiiy Callzptaminae. Below each penis apex (viewed from a postero-dorsal aspect except in B), is a transverse section through the valves to show relationships of penis and cingular vaives :

A. Indomerits Dirsh ; B. PaZaciosa C. Bolivar ; C. Peripolars Martinez ; D. Brachyxenia IGrby ; E and G-J. various a t present placed in Calopfenopsis 1. Bolivar ; F. Spho- dromevrcs Sta1 ; 1;. Paracaloptenirs 1. Bolivar ; L. Acorypha Krauss ; R1. Sphodronoius C‘varov. S and O. Inner apical surface of male cercus in C. coelesyriensis (Giglio-Tos) (S) and C. ikdiczcs (L.) (O). P and Q. Cerci of the genera Palaciosa C. Bolivar and Iizdomerzis Dirsh respectively. R-T. Apical abdominal segments of males of C. italicíts (L.), C. zunftemqliavus (Pantel), and C. bnrbarus (Costa) respectively. U. Inner pair of posterior tibial spurs in a species of Caloptenopsis 1. Bolivar showinp an intermediate stage in development of a pre-apical spur tooth (cf. Fig. 20: F and G which show estreme esamples).

29s S . D. JACO

-. Cercus not rugosely pitted on outer surface, relatively smooth, upper and lower mar- gins only slightly convergent apically or parallel. Penis valves developed or un- developed . 6

6. Apex of cercus trilobed, median lobe bearing an inivardly directed copulatory hook. If cercus bilobed then :

(i) outermost of inner pair of posterior tibial spurs unmodified, not strongly hirsute and with no pre-apical point (Test-fig. 20 : E. p. 324) ;

(ii) penis valves well developed being erect and as well developed as lateral accessory processes, not being merely an auricular fold of posterior inner edge of lateral accessory processes (Test-figs. 10 and z j . pp. 307 and 339) ;

(iii) lateral carinae of pronotum never strongly narrowed in anterior third. Lateral carinae of pronotum never obsolete, usually clearly defined ;

(iv) if cercus bilobed apically lower lobe never less than one third depth of upper lobe CALLIPTAMUS Serville

S -. Apes of cercus always bilobed . 7 7. Lateral accessory processes of each penis valve auricular on posterior face

that of cingular valve . . 11

. -. Lateral accessory processes exposing a fiat face posteriorly, their planes being same as

8. Lateral pronotal carinae obsolete. Posterior margin of pronotum very acutely produced . . BOSUMiA Ramme

9

dorso-lateral scales. Wings absent . . PARACALOPTENUS 1. Bolivar

-. Lateral pronotal carinae clear, or only partly obsolete . 9. Pronotum markedly tectiform. Posterior tibial spurs never modified. Teopina form

-. Pronotum not markedly tectiform. If tegmina reduced to scales then posterior

10. Upper inner area of posterior femur almost vertical. Dorsal carina, as seen from above, very close to inner side of femur. Disruptive band of light colour across outer surface of posterior femur, jus t proximal to knee, matching with a dark band across folded te,gmina, about 1/3 from their tips. Eyes light in colour, never mith pronounced eyestripes, though in life with structural iridescence indicating eyestripes. Frontal ridge above antennal sockets, widely expanded and flat .

-. Upper inner area of posterior femur clearly sloping ouhvards. Dorsal carina, as seen from above, not markedly close to inner side of femur, often lying on mid-line. No disruptive femoral band of light colour as described above. Eyes often dark monochromatic, or with pronounced eyestripes. Frontal ridge above antennal sockets slightly expanded, and usually concave

(There seems to be no satisfactory means of separating these n o genera) I I . Femur of second pair of legs with only upper furrow distinct on outer side ; loiver

carina of posterior femora widened beyond middle. Hind tibia with 6 (seldom 7) extemal spines . . SPHODROMERUS Sta1

-. Femur of second pair of legs with z distinct furroivs on outer side ; lomer carina of posterior femora not widened beyond middle. Hind tibia with g (seldom 8) extemal spines . SPHODRONOTUS Uvarov

tibial spur elongate and modified . . 10

STOBBEA Ramme

ACORYPHA Karuss and CALOPTEXOPSIS 1 . Bolivar

t

CALLIPTAiMUS Serville, 1831 Calliptaiiizrs Serville, 1831, d n n . Sci. nnt. (Zool.), 22 : 251. Type species Cnlliptarnus italicus

Caloptenzts Burmeister, 1838, Haiidbrick der Entomologie. 2 ( 2 ) : 637. Emendation for Callip-

Callipteniis Stal, 1S73, Recelisio Ortlioptevovii~~i. 1 : 3s. 72, 73. Emendation for Calliptamus

(Linnaeus, 1 7 j S ) . [Xeotype in British ,\Iuseuni.]

tantzis Serv., 1S31.

Serv., 1S31.

?

200-

180

160

140

120

loor

80

A RET-ISIOK O F T H E G E S U S C . 4 L L I P T A M U . S S E R V I L L E 299

.%fetromer¿rs Uvarov, 1935, Aitn. illag. zat. Hist. ( I 1) 1 : 3 j9 . T>pe species iifetromertrs coeles-vriensis

I . ( n ) Xedeagus composed of an upper cingular valve (Text-fig. I : a, and Text-fig. 10 : E and G, a. p. 307), andpairof penisvalves (Text-fig. I : m, and 10 : E-G, m. p. 307). Penk valves always with lateral accessory processes (Test-fig. I : 1 ) . (b) Cingular valve tongue- like with membranous lateral processes of penis valves (e.g. C. italicirs (L.), Test-fig. 10 : C, D, F. p. 307), or thickened and straight apically (Test-fig. 10 : E. p. 307, and Test-fig. 25 : C and F. p. 339) with penis valves which project b e o n d tip of cingular valve, their lateral accessory processes merely abutting on cingular valve laterally, not encircling it. Penis valves of latter type orientated in a plane at right angles to that of cingular valve, or if broad and lobate and lying in same plane as cingular valve, (Test-fig. 16 : B. p. 31 j) then cingular valve itself membranous (cf. Indoinenrs Dirsh, couplet 5 in key p. 297, and Text-fig. 3 : A. p. 296).

(c ) Penis valves either elongate, blade-like, and pointed (Text-fig. 10 : C, m. p. 307, and Text-fig. 13 : A, B, m. p. 311) or blunt and more or less thickened, with slightly to well developed flanges along their upper outer edges, and thickened rugose lateral accessary processes (Text-fig. 2 j : F, p. 339). Lateral accessory processes of latter type may be auricular, and shield weak penis valves (Text-fig. 15 : F. p. 315).

(Giglio-Tos, 1893). [Type in Turin 1Iuseum.i

DIAGNOSIS s.

-i

- c 3

.

-

-

-

X

X

X O C. cvrenoicus x C . b. barbarus X

6 0

Y . . . . 0 .

Fem. i.

eo 100 120 140

A scatter diagram for the ratio of femur length to wing length in males of C. barbnrrrs (Costa) nnd C. cyveitaicrrs sp. n., from various North African and European populations. C. bnrbnrzis bnrbnrrrs (Costa) from the Iberian peninsula, and C. cyrennictrs sp. n. show close affinities, while the scatter diagrams shift toivards higher values as one moves easbvards in southem Europe.

FIG. 4.

300 N . D . J A C O

2 . Cerci usually tri-lobed apically, upper lobe broad and laminar, lower lobes smaller, middle lobe bearing an inwardly directed hook (Text-fig. 3 : O. p. 296). Lower pair of lobes may however show progressive fusion giving a bi-lobed apex ; invariably so in C . coeiesyrien~is (Giglio-Tos) except h its subsp. hissa~icus (Mishchenko) (Test-fig. 17 : M. p. 319). 3. Inner pair of posterior tibia1 spurs unmodified, outer spur of pair never strongly elongate

or hirsute (Text-fig. 20 : E. p. 324). 4. Pronotal carinae distinct. Lateral carinae may fade just before reaching posterior edge of

pronotum (Text-fig. 19 : i3-J. p. 322). Pronotal sides almost vertical (cf. Bosirmza Ramme and Brachyxenia Kirby), except in upper quarter ; smooth whitish nodular area often present towards upper anterior comer (Text-fig. 20 : D. p. 324) (as in many Caloptenopsis spp.).

9. N O known diagnostic characters peculiar to this sex. Females very uniform morphologically throughout related genera in the sub-family. Cerci unmodified.

DiscUSSioN.-Synonymy of Metromerus Uvarov with Calliptamus Serville is

Fem.L. 12 14 16 18 20 22 24mm.

A graph of wing length against feniur length for females of C . trrriricercis Tarb., shoaing the regression line for points plotted (y = 1.15 x - 1.37, correlationcoefficient r = 0.7). Small circles represent material from Turkey (Ankara. hfersin, and Adana), U.S.S.R. (hzerbaydzhan, Daghestan, Tbilisi, and Yerevan), and Iran (Gandzha, TehrZn area) ; crosses represent material from Turkey (Urfa), Jordan (El Boweida, Wadi Zarqa. El Salt, and Shueib, and U.S.S.R. (Leninakan and Aresh). Large circles represent material from Israel (Negev), Jordan (Khor Kabid), Iran (Ahvaz), while solid dots represent material from Lebanon. Points with the lowest valuec for the ratio appear to belong to populations found in areas of highest rainfall (10 in. to 25 in. mean annual rainfall), the smallest coordinate values from areas with a mean annual rainfall 3s small as 4.9 in.

FIG. 5 .

A R E V I S I O X O F T H E G E N U S C A L L I P T A M U S S E R V I L L E

considered justifiedbecause the characters used to separate the genera areappiicable to either. Indeed Uvarov found the genus difficult to define when he revised it in 1943.

(i) Cercus morphology of JIetromerau Uvarov is closely approached by many specimens of C. sirbalpintts, C. zattenwylianzrs (Text-figure 24 : H, J, X.p. 336), C . turaniczis, and C. siciliae (Text-figure 24: XVIII. p. 336), as well as montane forms of C. barbarus (Text-fig. 24 : XI, XIV. p. 336). Moreover the subspecies C. coelesyriensis hissariciis often has a triiobed cercus apex.

(ii) The male genitalia show oniy specific differences, i.e. no more different than C. italiczis is from C. barbarus.

(Ui) Both C. watten-ylianzrs and C. coelesyriensis have melanic colour forms. (iv) The lateral pronotal carinae do not differ markedly from those of Calliptamtis,

a point indicated by Uvarov when he erected the genus. Examination of the type of Calliptamzts mus 1. Bolivar show that it was misidenti-

fied as a member of the genus Calliptamus and should be placed in the genus Sfihodymerzu St&l.

30 1

POLYMORPHICJI AND GEOCRAPHICAL VARIATION. The COmpOnentS Of POlY- morphism in the genus can be illustrated by five main characters :

FIG. 6 . The distribution in southern Europe and S o r t h Africa of C. ifnliczis (L.) (dotted and broken line), C. x-nftenwylinnrrs (Pnnt.) (obliqiie shnding) ; C. sz<bnlpinzts sp. n . and C. siciliue Rme. stat. n. (broken h e ) .

302 S . D . J . 4 G O

(i) Black spots on inner side of posterior femur. Three in number (Test-figure, 2: B, E, F, G. p. 293) but subject to various degrees of reduction (Test-figure 2 : H. p. 293), or expansion and fusion (Test-figure 2 : c, D. p. 293). Contraction in size, though not with a loss of intensity, takes place in populations subject to a n-et, cold climate. Expansion and fusion occurs under the opposite extreme in climatic conditions, namely dry and hot. Thus popdations from arid semi-desert may possess femora of the type shown in Text-figure 3 : D. In the case of specimens from cold dry environments, such as the uplands of centra1 han , the size of the middle spot may remain the same but the pigment becomes ven; diffuse, often only resdting in a greyness of the pink background colour, e.g. C. coelesyriettsis Izissnricus (Text- figure 17 : T. p. 319). C. barbnrus, and C. bulucha.

Especially well developed in C . wnttenwylinnzis is the tendency for a diffuse black pigment to be laid down throughout the pink or orange background colour of the postenor femur and tibia. This occurs in the south of its range of distribution in ail the drier, more and conditions of Korth Africa.

Occasionally where the melanic femoral spots disappear the black pigment can be replaced by pink, e.g. C. coelesyriensis (Text-figure 17 : P, Q). Apart from this facies, however, boreal and montane forms, living in wetter colder environments, have crimson as background colour. This may be deep or faint, the intensity of pigmentation being greatest at the extreme of coldnesc and wetness. Boreal species, e.g. C. nbbrecinttrs, will all have this facies ; those like C. italiczcs which enter more southerly environments wiil show

(U) Background colour of posterior femora and tibiae.

45

3 0

O 105 120 135 I I

FIG. 7 . C. tiirnniczcs Tarb., C. nbbrezintirs Ikonn., and C. itniicus (L.) : distribution in Middle East and Asia.

C

3

i

A R E Y I S I O S O F THE GESUS C A L L I P T - 4 J I C S SERYILLE 303

a fading of the red coloration, e.g. in Crete and the Greek Islands, and in Afghanistan, C. ita2ictrs may show little or no pink background pigment.

Incorporated with the red component, it produces an orange colour. Overlaid with a diffuse black pigment it becomes greyish green in appearance. C. tenzcicercis, C. bnrbaviis, and C. barbarus palnestiizensis all show the orange form, and in addition may also show yellow-legged forms in which the red colour component has disappeared (e.g. C. barbarus palaestinensis is entirely yeliow-legged, except in its abrupt entry into the Lebanon ranges where it becomes red-legged). Popuiations of C. bnvbnrits in southem Greece, e.g. Mt. Parnassos and Silókastron, show orange-red legs in the males while sympatnc females are yellow-legged. Also, while in Cyprus only one specimen of C. barbarus from hIt. Troodos was orange-legged (the rest being yellow-legged), mixed populationc of orange and yellow-legged inditiduals were usual in areas such as south eastern Turkey, and northern Syria (Text-figure 22 : A. p. 330). I t is quite clear that warm, dry conditions favour the formation of orange and yellow-legged polymorphs. The latter are general in areas affected by maritime conditions, such as the extreme north of hlorocco, or the Greek archipelago, or in cold dry iipland areas which experience very hot summer conditions, e.g. the Ankara plateau in Turkey, or central Israel.

In montane or extreme boreal populations, the hind uing pigmentation may be completely lost. This is fixed as a species character in C. nbbreviatus.

(iii) Tegminal length. híontane and boreal forms tend to be chort-winged, while the warm and often semi-desert conditions of the southern Palaearctic region, or inland steppes exposed to dry continental climates, show populations in which the wings and tegmina are weii developed, and often have apices which supass the knees of the posterior femora when the tegmina are folded.

In C. barbarus in Morocco, C. bulucha brnchyptertis in the Hindu Kush, and C. italiczrs in Afghanistan, tegmina may become so reduced as to be useless for flight. C. cyrennicirs, C. sttbnlpinzrs and C. abbrezlintzrs, are al1 speciec with tegmina of medium length, the apices of which never surpass the apicec of the posterior femora. C. italiczcs, C. tiwanictis, C. bnrbartis and C. teituicercis, are aii species showing every intermediate between elongate wings and tegmina iike those of C. subnlpinirs.

While members of the " ifnlictcs " group (see p. 320) tend to show iittle polymorphism in this respect (C. wattew~ylialzzts being a notable exception), the " barbnriis " group tends to show marked general body colour polymorphism. C. cyrenaicus, a species of the semi-desert fringe of Libya in North Afnca, show this exceptionally weii (see pronota, Text-figure 19 : B-J. p..322). The pronotum is not the only part of the exoskeleton to vary in colour, e.g. the head, Text-

respectivelJ-. 3iost specimene of C. italictu, C. nbbrevintirs, and northerly populations of other speciec, uill have pronotal types D-F. The uniform, pale coloured body colour, reprecented by pronota GJ, is found in semi-desert forms of C. itnlictrs and C. coelesyrieitsis, and many individuals from southerly populations of C. bnvbnrus and C. teniticevcis. The most estreme type of polymorph is f. iiinrginellics, repreeented by pronotum A, in which a dark, often brown body colour, is contrasted with 2 light cream ctripes on the vertes n-hich pass along the inner edge of each lateral pronotal

24§§

The antithesis to the red colouring is a yellow pigment.

(iv) General body colouring.

}

, figure 20 : A-C, p. 324, corresponding to pronotal types J, E, and F of Test-figure 19

E S T O M . f 3 , 9

304 K. D . J .4GO

canna, and may or may not be continued as a light line along veins Cit. and IA of the tegmina. Ulost memberc of the " itnliczis " group show this form to come degree, but although the northerly elements of the " barbnvtrs " group are often almost completely uniform in general body markings, southerly species and populations show the " inargiltellzrs " form as a dominant and striking polymorph. The various polymorphic forms grade into one another and are not clearly demarcated.

(v) Cercus apex. Only in C. coelesyriemis coelesyriensis is the bilobed cercus apex constant. In C. coelesyriensis hissariciis the middle and lower lobec are often distinct. In contrast, montane or boreal populations of C. baibnrzts, C. sitbnlpintts. C. wattenzylianacs, C. ttcraitictcs, C. bnliicha, and C. siciliae, chow a tendency for fusion of the lower pair of apical lobes, finaily producing a single lobe. Popuiations from semi-desert areas have a tendency to have clearly demarcated lobes.

When describing species, montane, boreal, or semi-desert forms may be referred to. Their characteristics can be deduced from above, if it is also remembered that boreal and montane forms are always smalier than semi-desert individuals of the same group.

There seems to be some evidence that the colour polymorphs and tegminal varia- biiity in Callipfamus may not be rigidy fixed genetically, but may be linked closely with the climatic environment. Thus the isoclines for the ratio femur length/tegminal

30' 350 4oo 450 50' 5 5 O 60' 65' 70'

The distribution in the Middle East, Turkey, and U.S.S.R. of C. coelesyriensis coelesyiensis (G.-T.) (broken h e ) , C. coelesyriensis hissnriczrs QIishch.) (northem obliquely shaded portion), C . baltrcka Uv. (southern obliquely shaded portion), and C. tenziicercis Tarb. (continuous line).

FIG. 8.

_-

FIG. 9. The distribution in Jordan rift valley area of C . barbarus barbarus (Costa) (vertical shading), C. barbarlis pnlaestinensis Rme stat. n. (diagonal shading), C. tenuicercis Tarb. (horizontal shading), and C. coeles*vriensis (G.-T.) (smail círcles).

a

3

I

i

35 36 37

306 S . D. J . A G 0

length inmalesof C. barbnrzts (Text-figure I I , ~ . 309) andfor the ratio femur length/head width in females of C. tentiicercis bear a striking paraiiel to the isohyets for the Middle East (Test-figure 31, p. 329). Femoral colour polymorphs also seem to be correlated with these isohyets. Thus C. temiicevcis, whose range seems confined within an area whose rainfall is less than 2 j in. per annum (cf. Test-figure g, p. 30 j, and Text-figure 21, p. 329), produces the colour variety f . niirnnti$m (pale orange tibiae) in the higher rainfall areas it enters in southern Turkey and western Syria. Comparison of C. barbavtis and C. tenziicercis leg colour polyrnorph distribution (Text-figures 27 : A, and 22 : B, p. 330) with Test-figure 21, shows that the orange femoral coloration is confined to areas with a rainfall of less than 2 j in. per annum in the former species, and less than 15 in. in the latter. The two species therefore appear to react to the enmronment differently, the orange-legged facies in C. tentiicercis being l e s tolerant to increased rainfall than that of C. bnrbnvzrs. If the wing length is plotted against femur length in females of C. tenziicercis (Test-figure j, p. 300), a regression line can be drawn (suggesting continuity of this vanant across the apparent diccon- tinuity of the leg colour variants), the values at the lower end of the graph corresponding to the areas of highest rainfall (10 in.-25 in. mean annual rainfaii per annum), those at the upper end to areas with low rainfall (often as small as 4.9 in.). Unfortunately the climatic data available refer to macroclimatic conditionc, so that conclusions must be tentative. I t is also ve" unlikely that the vanants described w d l correlate cimply with any one climatic factor.

KEY TO SPECIES MALES

I. hlembrane covering penis valves estemally produced into a long, decurved, backwardly directed pocket, i.e. with same configuration as penis valves inside (Text-fig. 3 : R. p. 296 and Test-fig. 10 : C, D, and F. p. 307). Hind wings never colourless, even if reduced . . itolicus (L.) (p. 316)

-. Pocket covering penis vavles not so elongated or shaped. Vsually bluntly produced and backwardly directed (Test-fig. 3 : T. p. 296), but ma? be erect, short, and

2. Inner face of posterior femora without any markings between upper and lower inner carinae ; this area pale body colour, without any trace of pink or orange pigment

-. -4 single black spot, or hvo t o three separate spots, between upper and lower inner

3. Pocket covering penis valves with outline shown in Test-fig. 3 : S. p. 206, orientation being vertically or slightly fonvardly directed. Only occasionally with median hooked apical cercus lobe wholly fused with ventral lobe, but always rather weakly developed. Rami of cingulum convergent ventrally (Test-fig. 13 : B. p. 311). Male phallic comples, Test-fig. 13 : A, F. p. 31 I

-. Pocket covering penis valves bluntly produced in a postero-dorsal direction (Test-fig. 3 : T. p. 296). Cerci invariably with median lobe fused into lower lobe, upper and lon.er lobes thus being almost equal to each other (Test-fig. 3 : S. p. 2gh, and Test-tig. 17 : L, S. and O). 31ale phallic comples Test-fig. 16. p. 317

pointed (Test-fig. 3 : S. p. 296) . 2

3

carinae of posterior femora . 4

. .turanicus Tarbinsky (p. 325)

coelesyriensis (Ciglio-Tos) (p. 343) 4. Posterior femora yellow 0x1 inner face . 5 -. Posterior femora dull crimson, orange, or red, betweeii upper and lower carinae, inten-

sity of colour varying from pale to deep . 6

. . .

I

'. .

:

A R E V I S I O S OF THE GESUS C A L L I P T A J I C S S E R V I L L E 307

I b

A

0.5 mm.

9 B D

b

/

E F G

FIG. 10. A and C . Lateral aspects of penis valve in C. nObver.intrts Ikonnikov and C . itaZicrrs (L.) respectively ; B and D. Dorsal aspects of penis valves in C. abbreviatus Ikonnikov and C. i tdici is (L.) respectively ; E-G. Transverse sections through valves of penis in C. bnrbnviis (Costa), C . italiczrs (L.), and C. rcaltenzqliatzirs (Pantel) respectively .

30s S . D . J .4GO

5 . Inner side of Dosterior femora filled or Dartlv filled. between umer and lower inner *.

carinae, by a solid black blotch, or separate spots which show various degrees of fusion (anterior pair first-as in Test-fig. 2 : B-D). Lateral appendices of penis valves bluntly pointed as seen from above ; valves strongly developed though may be pointed with triangular dorso-lateral espansions (Test-fig. zj :, A-D, F, J. P. 339) . . barbarus (Costa) (p. 327)

Inner side of posterior femora normally with 3 separate black spots but occasionally, when sympatric with orange-legged form of barbarirs (Costa), has a single solid inner femoral blotch. Lateral appendices of penis valves auricular. Penis valves weak (Text-fig. 1 5 : F, m. p. 315) . . tenuicercis Tarbinsky (p. 340)

Cercus \&h very large-laminar dorsal apical lobe (Text-fig. 2 4 : S I S . p. 336) ; lower lobes weak and divergent. \Vings smoky. hlale phallus, Text-fig. 26 : A, B, C. p. 346 . . piebeius (Walker) (p,

Cercus without such a large laminar dorsal lobe ; lower lobes may be fused but not attenuate .

Hind wings usually pink; only colourless in forms lvith very short tegmina from montane localities, in xhich wings no longer functional for flight .

Hind wings colourless, when folded not surpassing knees of posterior femora, but fully functional for flight .

Cerci -4th lower apical lobes separate ; median lobe distinctly longer than lower lobe. Male phallus, Text-fig. 10 : -4, B. p. 30 j . .

Cerci with median apical lobe more or less fused to lower lobe, equal to it in length or even shorter (Text-fig. 24 : XVII. p. 336) .

IVidest part of penis valves (as seen from above) clear of posterior edge of cingular valve. Cicil- only (Test-fig. 24 : XVIII) . . . siciliae Ramme (p.

Widest part of penis valves (as seen from above) leve1 with posterior edge of cingular valve (Test-fig. z j : E. p. 339) . . subaipinus sp. n. (p.

Inner side of posterior femora orange. Inner femoral blotch filling, or almost fillhg, inner area .

Inner side of posterior femora red, scarlet, or dark ruby ; varying intensity of colour. Inner femoral spots fused or separate

Posterior end of inner femoral spot showing at least come trace of orange a t its margh

Posterior end of inner femoral blotch without any trace of orange a t its margiii

Three separate inner femoral spots (sometimes pale) Anterior spots on inner side of femora fused, or one blotch present of smaller or larger

size, diffuse or clearly demarcated . F'ertical knob-like thickening on posterior median edge of dorsal ectophallic plate.

Penis valves typically lying in same plane as cingular valve. Cerci usually bilobed apically (Test-fig. 16 : C, D. p. 317) . . coelesyriensis (Giglio-Tos) (p.

Posterior margin of dorsal ectophallic plate laminar (Test-fig. I, p. 291). Cerci variable apically .

inner side of posterior femora with large diffuse median blotch, general colour sur- roundiiig it being sniokey ruby-red (appearing dull mauve). Tendency to be moderately or v e n brach-pterous (then non-functional for flight). Folded tegmina with apices never surpassing knees of posterior feniora. Cerci less than 3.j times longer than broad (see ttnuicevris Tarbinsky, couplets 5 and I 1, where cerci at least 4 times longer than broad) . . balucha Uvarov (p.

Inner side of posterior femora, if approaching colour described above, with 3 separate spots of roughly equal size. Sever with non-functional hind wings

Aedeagus with pointed, non-auricular, lateral accessory penis valve processes (Test- fig. 13 : 6, C. p. 311) curling upwards and lying above cingular valve. Pocket

. abbreubtus Ikonnikov (p.

.

barborus (Costa) (p.

tenuicercis Tarbinsky (p. .

.

347)

7

1 0

8

326)

9

340)

338)

11

12

327)

340) 13

20

343)

14

342)

15

i

c

-4 R E Y I S I O S O F T H E GENUS C A L L Z P T A i I I U S S E R V I L L E 309

FIG. 11. C. barbarzrs (Costa) ; isoclines of femur length to tegminal length ratio for males from Middle East and Central Asia.

b

FIG. 12. C. teiziricevcis Tarb.; isoclines of femur length to head width ratio for females from hliddle East and Central Asia.

310 X . D . J A C O

16. Inner side of posterior femora dull pale pink ; tibiae pale to dark orange-yellow

-. Inner side of posterior femora dull ruby-red, posterior tibiae crimson or dull ruby-red 17. Folded wings with apices not reaching level of tips of posterior femora. Xedian and

lower lobes of cercus tip always intimately fused to each other, median lobe equal to or even shorter than lower . 18

-. Wings not always as short as above, often when folded with apices surpassing knees of posterior femora. Median and lower lobes of cercus apex usually unfused, median lobe longer than ventral lobe. Penis valves boldly sclerotized, not greatly tapered or out-curved apically (Test-fig. z j : A-D and G. p. 339)

18. Penis valves small, usually tapered and slightly outcur\*ed apically. SE. France and Italian peninsula . . subafpinus sp. n. (p. 338)

-. Penis valves small, roundly truncate apically. S o t noticeably outcurved (Text-fig. zj : K. p. 339). Sicily . . sicifiae Ramme (p. 340)

19. Cingular valve, as seen from above, estending well beyond a line drawn between apices of lateral appendices of penis valves (Test-fig. 2 j : G. p. 339). Folded tegmina with apices never surpassing knees of posterior femora, and tapered apically. Cyrenaica . . cyrenaicus sp. n. (p. 335)

-. Cingular valve apex, as seen from above, level with a line drawn between apices of lateral appendices of penis valves (Text-fig. 20 : -4 and B. p. 324) -4pices of folded tegmina often surpass knees of posterior femora. (Never sympatric with cyvenaiczrs escept as orange-legged form with solid inner black femoral markings)

tenuicercis f. aurnntipes Ramme (p. 306) 17

19 .

barbarus (Costa) (p. 327) 20. Inner femoral spots fused to form a single blotch lying between upper and lower inner

carinae . 21 -. Only anterior pair of spots fused . . barbarus (Costa) (p. 327) 21, Lateral carinae of pronotum without any trace of convergence in metazone. BIale

penis valves weak, not extending far beyond cingular valve, as viewed dorsally. Lateral appendices of penis valves rather auricular (reminiscent of fenuicercis), whole structure recalling a large C. szrbal~inus (Text-fig. 26 : D. p. 346). Cingulum, a t its dorso-posterior edge, forming 2 gently down-sloping lobes madeirae Uvarov (p. 347)

-. Lateral carinae of pronotum usually convergent in metazone. Jiale penis valves strongly sclerotized, estending well beyond cingular valve as seen from above. Lateral eqansions of penis valves not auricular. Cingulum vertical behind its dorso-posterior edge . . barbarus (Costa) (p. 327)

FEJIALES (Provisional key)

Because of the morphological uniformity among females in this genus, the following key has been sub-divided geographically as follows : A. Southern Europe (escluding G.S.S.R.) B. Sorth Africa C. Turkey, G.S.S.R., the BIiddle and Far East.

X. Couthern Europe (escluding U.C.S.R.)

I . Hind wings colourless . 7 -. Hind wings infused with pink . 2 2 . Inner side of posterior femora pale body colour or yellow -. Posterior femora red, orange-red, or dull ruby-red on inner side

. 3 ‘ 4 .

311 -4 R E V I S I O S O F T H E G E S U S C A L L Z P T A A I U S S E R V I L L E

9

I

\

B a

FIG. 13. A. and B. Postero-dorsal aspect of phallic complex of C . tirvaizicirs Tarbinsky and C. zuattenzqylinnzrs (Pantel) respectively : C. Dorsal aspect of aedeagal apes in C. wattenwylinnus (Pantel) : D. Lateral aspect of whole of phallic complex in C . wuttcn- wylianzrs (Pantel): E and F. Dorsal ectophallic plate of C. wnt/erzwyeiylinrzzts (Pantel) and C. tztvanicirs Tnrbinsky respectively. (The arrows in X and B indicate upward diver- gente or upward convergence of the cingular rami in the phallic complex of each cpecies, this being one of the main diagnostic characters.)

*4§§§ ENTOM. 13, 9

3x2 S . D. J A G O

3. Three separate spots on inner side of femora. Spots often faint or almost absent from inter-carinal area (Text-fig. 20 : H and IC). Tegminal apices neier surpassing knees of folded posterior femora. Tegmina usually markedly tapered apically

-. Single black femoral blotch between upper and lower inner carinae of posterior femora (Test-fig. z : D. p. 293). Tegminal apices usually surpassing knees of folded posterior feniora, or leve1 with them. Tegmina not markedly tapered apically

tvattenwyfianus (Pantel) (p. 320)

barbarus (Costa) [p. 327) 4 . Tegminal apices clearly surpassed by knees of folded posterior femora. Tegmina

-. Tegminal apices surpassing knees of folded posterior femora, or if falling short of

5 . Large forms (total length 21.6-41.9 mm.). Distribution southern and eastern Spain,

tapered in apical 213 . 5

iatter, then doing so by only a very small margin

southern France. Kot occurring east of French JIaritime Alps.

. 6

wattenwyíianus (Pantel) (p. 320) -. Small forms (total length 20*6-28*6 mm.). Distribution Italy, from Ligurian Alps to

6. Inner femoral spots separate. Usually equal in size, often faintly developed, only just crossing upper inner carina (Text-fig. 2 : E. p. 293)

-. Median inner femoral spot usually bigger than other two. If faintly pigmented however, very similar to more darkly pigmented forms of ifulicus. Separable in such cases only by a study of males from same locality (ifuliczrs has longer tegmina) . barbarus (Costa) (p. 327)

7. Range in France and Italian peninsula . . subalpinus sp. n. (p, 338) -. Sicilian species only . . sicifiae Ramme (p. 340)

Sicily ; also in SE. France Plaritime Alps) . 7

. . italicus (L.) (p. 316)

B. Xorth Africa

I . \Vings and tegmina ver). short, never extending further than middle of folded posterior femora . barbarus (Costa) (p. 327)

-. Tegminal apices surpassing middle of folded posterior femora

-. Inner side of posterior femora red, dull ruby-red, or greyish pink .

. 2

2. Inner side of hind femora yellow or orange with black markings. Three black spots, sometimes fused . . barbarus (Costa) (p. 327)

3 3. Bulky insects. Colours not contrasting. JIelanic forms common. Tegmina often

markedly tapered in apical 2/3. Head 5'0-7'1 rnm., femur length iÓ*0-2j*0 mm., tegminal length 16'3-31.5 mm. . . wattenwylianus (Pantel) (p. 320)

-. h'ot conspicuously bulky. Often brightly coloured with " margirzdilirs " form in evidence. S o melanic forms. Tegmina only markedly tapered in apical 2/3 in cyrenuiczrs sp. n. Head width 4.756'0 mm., femur length 1q5-19.2 mm., tegminal length x y o - - 71.0 mm. . barbarus (Costa) (p. 327) or cyrenaicus sp. n. (p. 335)

(Separable on distribution or by males)

C . Middle East, U.S.S.R., Far East

I. Hind wings colourless, when folded never surpassing knees of folded posterior femora abbreviatus Ikonnikov (p. 326)

-. Hind wings pink. Often mhen folded surpassing knees of folded posterior femora

-. Inner side of posterior femora with a t least one darker or blackish marking

. 2 2. Inner side of posterior femora unmarked between upper and lower inner cannae, this

area being dull pink or body colour . 3 . 5

bodycolour . . turanicus Tarbinsky (p. 325) 3. Large insects. Head width 6-3-7.1 mm.; inner side of posterior femora dull pink or

i

-i

- .

-4 R E V I S I O S O F T H E G E S U S C . i L L I P T . 4 M C . S S E R V I L L E 3'3

F I rnm

1

5

FIG. 14. Epiphalli : A, B, D, and G. C . coelesyriensis (Giglio-Tos) from : -1. U.S.C.R., Armeniya S.C.R., pr. JIegri ; B. Turkey. Isparta prov., Dedegol Dag mt.. Sütcüler ; C. Iran, Haftom prov.. Lár ; G. (C. coelesyriensis hissnricrcs (Jlishchenko)) Afghanistan, Badakhshan prov., Cenna, 1,Soo m.; C. C. balucha Uvarov from West Paki- stan, Chitral reg., Chitral, 2,300 m.; E and H. C. uatte?zwyliantrs (Pantel) from E. France, Pyrénées-Orientales, intermediate to type found in Korth Afnca. H. France, Var, tq-pical of type found in south-eastern France. F. C. trtvaniczrs Tarbinsky from : U.S.S.R., Kazakhstan S.C.R., Yuzhno-Kazakhstanskaya Obl.

314 S . D. J A G O

-. Smaller insects. Head width 4*0-6~0 mm. . 4 4. Tegminal apices falling well short of knees of folded posterior femora. Tegmina

speckled with sniall flecks of darker colour. Inner side of posterior femora dull grey-pink . . coelesyriensis (Giglio-Tos) (p. 343)

-. Tegminal apices leve1 with knees of folded posterior femora, or surpassing them. Tegmina immaculate. Inner side of posterior femora pale body colour

italicus (L.) (p. 316) (buff desert form)

5 . Posterior tibiae yellow or orange . 6 -. Posterior tibiae red, or crimson, or dull pink . . 11

G. Tibiae orange 7 -. Tibiae yellow 8 j . Solid inner femoral blotch on posterior femora (ofteri estending below lower inner

carina) with orange pigment just posterior to it . . barbarus (Costa) (p. 327) -. As above, but blotch may be faint So trace of orange pigment posterior to black

mark or blotch . . tenuicercis Tarbinsky (p. 340) S. Inner side of posterior femora with solid inner blotch

tenuicercis Tarbinsky (p. 340) or barbarus-(Costa) (p. 327) (In southem Dead Sea valley and h-egeb desert : divided on distribution, i.e. Test-fig. g (p. 305), or by association with males.)

-. Inner femoral spots with anterior pair fused, or al1 separate, or clear evidence (if in one continuous irregular blotch) of tripartite origin . 9

g. Al1 inner femoral spots clearly separated . . 1 0 -. At least anterior pair of spots fused

tenuicercis Tarbinsky (p. 340) and barbarus (Costa) (p. 32 j ) (Inseparable escept on distribution, i.e. tenziicercis with these facies in northern S j n a and SE. Turkey ; barbarlis in Israel and Transjordan ; or by association with males.)

10. Head width 3'9-5.0 mm. Small insects. Central and southern Turkey tenuicercis Tarbinsky (p. 340)

-. Head width 5-0-6.4 mm. Larger insects. Israel and western Lebanon barbarus (Costa) (p. 327)

I I. Inner side of posterior femora dull greyish pink. Inner femoral spot thus diffuse. Often showing extreme brachypterism . . baíucha Uvarov (p. 342)

-. Inner side of posterior femora with three clearly separate spots, though sometimes faint. Feniora and tibiae never dull greyish pink, nor with large diffuse inner spot.

12. Conipletely black insects . . coelesyriensis (Giglio-Tos) (p. 343)

13. Te,mina creamy buff. Very finely peppered with small dark spots. Inner side of

So t showing estreme brachypterism . . 12

--. Buff, brownish, or greyish brown spotted insects . 13

femora with three spots, often faint (Text-fig. I 7 : S. p. 319) coelesyriensis (Giglio-Tos) (p. 343)

-. Tegmina dark brownish grey. Coarse spots or markings often orientated into trans-

14. Inner femoral spots estending only a short way below upper inner carina of posterior femora. Hind legs dull pink or dull mauve. \Vings and tegmina w l l developed (escept in Afghanistan where populations with short tegmina occur, i.e. knees of folded posterior femora surpass apices of folded tegmina, latter tapering in apical 2 /3)

verse bands across them . . 14

italicus (L.) (p. 316) -. Inner femoral spots larger and clearly defined, estending at lcast to middle of inner

15. Posterior tibiae and inner side of femora dull to bright crimson, though this colour may fenioral area. Hind legs dull mauve to bright crimson . . I j

disappear leaving almost al1 inner femoral area same colour as body. barbarus (Costa) (p. 327)

L 4

A R E V I S I O X O F T H E G E X V S c d L L I P T - d - \ r u s S E R V I L L E 315

D H m

0

m I

1 9 C

O K L

I m m

FIG. 15. Phallic complex of C. feitzticercis Tarbinsky, (D-F., K-U., and O-Q.), and C. balzrchn Uvarov (a-c., G-J, and S ) . (a) C. fei~tticeucis Tarbinsky : D. Lateral aspect entire male phallic complex (material from Turkey, near Ankara). E. and F. Dorsal and posterior aspects respectively of apes of vnlves in D. I í . Dorsal ectophallic plate. L-31. and O-Q. Epiphalli from : L. Turkey, .lnknra. 31, O, and Q. Jordan, El Ghor valley, Jerash Rd., Wadi Zarqa. (b) C. b a l r d n Uvarov : A-C. Dorsal, posterior and lateral aspects of tip of penis in subspecies braciij~pfrrirs (Dirsh) from Chitral. G- J. Dorsal, posterior, and lateral aspects of tip of penis in noniinate subspecies from West Pakistan, Baluchistan reg., Ziarat. X. Epiphallus of nominate subspecies.

316 S . D . J A G O

(Lebanon, S. and \V. Turkey, Iranian Azerbaidjan, U.S.S.R. bordering Black Sea, S . Caspian, etc.)

-. Posterior tibiae and inner side of femora mauve or dull pale ruby. Colour not intense coelesytiensis (Giglio-Tos) (p. 343)

(Turkish upland form)

Calliptamus italicus (Linnaeus, 1758) Gryllris Loczista italiciis Linnaeus, 1758, Systenta A-attirae, 10th edn.: 432. [Seotype 3, Italy,

Basilicata reg., Venosa, 420 m., 30 . i s . 1937, (F. E . Zeziner). Brit. Mus. (nat. Hist.) (subject to ratification by the Intemational Commission on Zoological Nomenclature)].

Calliptamzis itdiczrs grandis Ramme, 1927, Eos, 3 : 166. [Holotype 9, Italy, Sicilia I., Fontana hlurata, 17 .vii. 1924, (Ramme-Richter).] Syn. n.

Calliptamirs italicus reditctics Ramme, 1930. Mitt. zool. Mz<s. Berl., 16 : 214. [Holotype J, U.S.S.R., Kazakhstan S.S.R., Turkestan, Kazamuk Alai, SSjO m., 5 .viii. 1889, (Conrudt).] Syn. n.

Calliptamzrs italiczis inszilaris Ramme, i g j i , Mitt. zool. fifars. Berl., 27 : [Holotype J, Greece, Páros 1.. viii. 1925 (A . Sch~iltz).] Syn. n.

Calliptamus afghanus Ramme, 1952, Vidensk. Medd. daiisk Xatrtvk. Foven. Kbh., 114 : 200 [Holotype 8, Afghanistan, Faran and Chalchansur prov., Farah, 18 .vi. 1948, (Haadov, N. ) . ] Syn. n.

[Last four holotypes in Zoologisches ivfuseum of Humbolt-Universitat, Berlin.]

DIAGXOSIS d. I . Aedeagus (Text-fig. 10: C, D. p. 307), long and decurved. Penis valves elongate and blade-like. Lateral processes of penis valves (Test-fig. 10 : D, 1. p. 307) with their pointed apices curling up and over bases of decurved penis valves to form accessory stylets. Genitalia show greatest affinity with C. abbreviatus, C . wattenwylianus; and C . iztranicus, differing from the first in having penis valves almost twice as long [cf. Test-fig. 10 : A and B. p. 307), and from last two in not having cingular valve, and lateral accessory processes of penis valves roughly same length (Test-fig. 13 : C and D. p. 311). Asis of aedeagus directed, in situ, upwards and backwards, valves enclosed by digitiform pocket in ectophallic membrane (Text-fig. 3 : S. p. 296). Cingular valve membranous, elongate, intimately attached to dorsally curled parts of lateral accessory valves (Test-fig. 10 : D, a. p.307 ). Posterior margin of dorsal ectophallic membrane plate laminar. 2. Cercus usually with al1 three apical lobes well developed and unfused (like Text-fig. I 7 :

H. p. 319). Cercus like that of many C. barbarus and C. tenuicercis. 3. Tegmina usually well developed, when folded estending beyond knees of folded posterior

femora. Occasionally abrupt shortening of tegmina occurs, tegmina then tapering in their apical 2 /3 (montane forms) .

4. General coloration dark, escept in very dr?;, hot regions, where light buff specimens occur (pronotal type G , Test-fig. 19. p. 322). Pronotum usually similar to F. Test-fig. 19. p. 322. Colour variation less than C . barbarzis ; extreme " marginellzis " variety never occurs (Text-fig. 19 : B. p. 322 ; and p. 303).

5 . Posterior femora intemally with 3 dull brown or blackish spots which are alwa)rs separate (Test-fig. 2 : E. p. 293). Median and posterior spots equal in size, often estending only a short way below upper inner carina of femur. Confusable w t h C . barbarus in montane areas, since barbariis may have similar reduction and concentration of femoral spots (see Test-fig. z : H. p. 293). Populations of dry steppe often have almost immaculate pale creani inner femoral area (like C . tziraniczis).

6. Posterior tibiae invariably some shade of dull ruby-red, black pigment sometimes giving a mauve effect. Inner side of feniora usually weakly pigmeiited with pink along and adjacent to low-er inner carina.

308.

7 . Hind wings always with some trace of pink spreading across anal fan irom wing base.

t

3'7 A R E V I S I O S OF T H E CESC'S C A L L I P T . 4 . V U S S E R V I L L E

C

rn

I H J

F I mrn 'd G

Fxc. 16. Phallic comples of C . c o e l e s p i e m i s (Giglio-Tos) : A , C , E, and G. from Iran, Lar-ty-pical of semi-desert populations. A. Dorsal aspect of tip of phallic comples shoiving penis valves. C. Lateral aspect of entire phallic comples. E. Posterior aspect of apex of phallic comples. G. Dorsal ectophallic membrane plate. B, D, H, J , and F. Phallic comples of C . coelzsyiem-is hissnvicris (Jfishchenko) : B, D, and F. Posterior, lateral, and dorsal aspects of apical part of penis comples in material from hfghanistan, Badakhshan reg., Senria. H and J. Posterior aspects of intermediate penis types from zone of contact with nominate siibspecies. Iran, Dovvom prov.. Shahrud.

.. . ' ,,

3 1 s P;. D . JACO

9. Often very difficult to differentiate from C. barbarus, C . watte)tw~iianzcs, and C. srrbnipinus (sympatric with these spp. in SE. France), but in such localities uauallv having long wings (when folded har-e apices leve1 witli or surpassing knees of posterior !emora), and subequal, separate, inner femoral spots which fade ventrally (in C. bnrbarzis median of 3 spots usually largest, i.e. Test-fig. 2 : E-D, F-H. p. 293) .

~ L E A S U R E M E X T S Males Females - -7 Head width . 2 1 3 3 .4-4 .6 3 .83 . 3úS 4 '4-6.2 5 13

KO. Range JIean S O . Range Mean

Femur length . 213 9'7-14'7 1 1 . 5 0 . 370 13.8-24.6 17 .65 Tegminallength . 214 10.4-'2.? 1 j . 3 7 . 364 14.2-31.3 ~ 3 . ~ ~ Total length . 217 16.1-28.7 21 .09 . 371 21.9-41.G 3 2 - 1 0

DISTRIBUTIOX. So t found in Sorth Africa or Spain south of the Pyrenees, it extends across southem and south-eastem Europe (see Text-figure 6, p. ~ o I ) , western and southern Turkey, and follows a belt of cooler conditions into central Asia (Text-figure 7, p. 302). Its populations overlap in the east with those of C. nbbreviatus. The species has the most northerly distribution of any species in Europe, extending down the Rhine following drier " Mediterranean " conditions.

XATERIAL EXAJIIXED. FRANCE : Pyrénées-Orientales, 8 $, 1.2 $2, viii-xi ; Hérault, 5 8, 3 9, viii ; Paris, 2 3, - ; Seine-et-Mame, I 8, vii ; Haute Loire, 4 3, 3 Q, vii ; Puy-de-Ddme, I 8, 3 9, vii ; Bouches-du-Rhhe, I 8, vii ; Lozere, I 3' vii ; Alpes Mantimes, up to 660 m., 19 3, 41 $2, viii-x ; Var, up to 3jo m., S G, 14 Q, viii-x ; Corsica, 2 3,4 Q, vii-ix. SWITZERLASD : T'alais, up to 2,000 m., I 2, viii. AUSTRIA : Bwgedand, nr. Iilmitz, I Q, viii ; Salzburg, 5 d, viii ; L. Austria, I 9, viii. GERMANY : Hescen Iander, I 8, I 9, viii ; Bayern lander, I 2, viii. ITALY : Spezia, I 3, I Q, vii ; Ravenna, 2 Q, vii ; Basilicata, 420 m., 2 d, ix ; Grosseto, 60 m., I 3 ; Puglia, 20 m., 2 9, ix ; Sardinia, I 8, 3 Q, vii. JUGOSLAVIA : Hrvatska, 10 9, 9 9, vi-is ; Srbija, 680-1400 m., 9 2, 1.2 9, viii-s ; Crna Gora, 14 2, 17 9, viii-ix ;

c

FIG. 17. -1.-D. Fastigium verticis of 3 males (A-C) and a female (Di of C. coelesyriensis coelesyriensis (Giglio-Tos) from a single Turiiish population, to show range of variation. E-K. JIale cerci : E and F. Lateral and dorsal aspects in C. bnlzrclza balirchn Uv. G. and K. Lateral and dorsal aspects in C . balzrcha bvnchypferzrs (Dirsh). H and J. Lateral and dorsal aspects of cerci in C . teizuicercis Tarbinsky populations from iir. Amman, Jordan. L-O. Outer surface of npes of right male cercus in C. coelesyriemis (Giglio-Tos): L. Sominate subspecies, Turkey, fsparta prov., Sütcüler. Ji. Subspecies hissnricits (JIishch.), Iran, Shahrud. S. Sominate subspecies, Turkey, Mugia prov., Sandras Da@, 1,500 m., (intermediates betneen tn-o subspecies). O. Sominate subspecies, Iran, Haftom prov. Lar (typical of larger specimens from semi-desert areas). P-U. Inner aspect of left posterior femur in C. coe izsp ie ix ic (Giglio-Tos) sliowing estent of blnck markings and intensity of pink pigmentation : P-R nnd L. Males ; S and T. Females. Al1 nominate subspecies unless otherwise stated. P and Q. Turkey, 3iu&-1 prov., Sandras Da& I,SOO m. R. Turkey, Crfa prov., TIrfa (semi-desert non-melanic form). S. Turkey, Antalya prov., Gebiz (dark plateau form). T. Iran, Dovvom prov., Shahriid (subspecies hissaticzis (Nishch.)), pinli area being suffused with black pigment giving a dull mauve effect. U. Iran, Dowom prov., Shahrud Oight semi-desert form). In al1 femoral diagrams, backgroiind colour pale buff or straw colour (same as abdomen).

A B C D

E H

i*,rnrn- J

.._. .,. ............... .' .. :... c- : . .. :*"i,,, -...........,A ...... c .......... L

N

K

.... ...... O

block pigment 1 pink pigment

320 S . D. J a c o Vojvodina, 10 g, 21 9, vi-viii ; Hercegovina, I a, I ?, is . ALBAXIA : Gjinokaster, I 9, 3 Q, viii-ix. GREECE : Makedhonia, 17 Z, I Z 9, vi-Liii ; Kefallinía I., 7 3, 4 Q, vii ; Évvoia I., 2,100-2,600 m., 14 3,12 G, Stereá Ellás, up to 1600 m., I 3, 3 9, vii ; Icérkira I., 4 8, 5 9, vii ; Trikkala, 2,000-2,300 m., I Z, vii ; Samothráki I., I 8, I 9, vi ; Kriti I., up to 1100 m., I 3, 6 9, vii-viii. CYPRC'S : Limassol, 2 9, 2 9, v. TL'RKEI- : Antalya, up to 1700 m., I d, ij 9, vii ; Istanbul, 9 a, 11 $2, vii ; Kocaeli, 2 ,j', 29, viii; Izmir, I d, 2 Q, vii ; Mugla, 2 Z ,4 9, vii ; Konya, 2 Q,- ; Sinop, I 9, ix ; Urfa, 2 3, I 9, vii ; Isparta, 1,600-1,700 m., 2 Q, Mi. IRAN : Dovvom, 4 9, 14 Q, ti-s; Hashtom, I 3, t- : Dahom, 2 0, 3 9, vii-ix; blarkazi, I d, 2 Q, vii-viii ; Xohom, 7 8, 2 Q, vi. AFGHAXISTAS : Mazar-i-Sharif, 5 d , 4 9, vi ; Badakhshan, 1,S00-2,300 m., 5 3, 17 Q, vii-viii ; Kataghan, 625-1,240 m., I o', 2 9, ix ; Kabul, 1,600-1,740 m., 7 9, is-x; Eactem Suristan, 2,700 m., I $?, vii. U.S.S.R. : Kazakhstan S.S.R., 12 d, IOI Q, vi-is; Armeniya S.S.R., 3 g, 13 Q, vi-viii; Daghestan A.S.S.R., 8 8, 4 Q, viii-ix ; Tadzhikistan S.S.R., I d, vii.

DISCUSSION. Aithough capable of living in relatively cold, moist environmentc, this species cannot tolerate wet aipine conditions. It does, however, penetrate into dry alpine conditions up the R h h e vaiiey to Lake Geneva. It is the dominant species of the cold steppes of southern U.S.S.R. in the regions of the Black Sea, Cacpian Sea, and Aral Sea, but in Southern Europe takec second place to C. barbarus. Amongst hundreds of Cnlliptamus specimens from Cyprus, ody 5 were C. italicus. Unlike C. barborus it doec not thrive in estreme dune conditions, e.g. Rh6ne delta, and never reachec habitat altitudes like those of C. strbal$ifizis (p. 338). La Greca (195s) statec that in Italy the species occurs from sea leve1 up to 1,000 m. It cannot tolerate the extreme aridity enjoyed by C. bnrbnrzts and therefore does not extend so far south as this speciec. None of the subspecies claimed for this insect seem to be valid. The male phallic complex shows complete uniformity throughout the range of the cpecies, and other variations can be attributed to environmental effects (Discussion, pp. 301-306). The species shom little colour polymorphism except on the extreme southem edge of its dictribution, C. bnrbnriis displaying marked colour polymorphism even when sympatric with uniform C. itnliciis populations.

The only specimen of Calliptnntzis in the Linnaean collection is unlabelled, and is a female of C. watte.nwylia.ntis. Dr. -4kZ Holm informs me that there are no cpecimenc of C. itnlicirs labelled by Linnaeus in the Uppsala collections. 1 am applying, therefore, to the Intemational Commission on Zoological Xomenclature for formal authority to reject the Linnaean specimen as type of Grylliis (Lociista) italicus and to designate a neotype. ,4n application has been sent to the Secretary.

v:

Calliptamus watteniuylianus (Pantel, ISgG) Cnlopfe i im italicris var. wattentiylianiis Pantel, 1596, A n . Sor. esp . Hist. mt., 25 : 70, I pl.

:Holot?pe o', Spain, RIalaga prov.. Sitio. Paris híuseum.] Calopfeitirs okbaensis Kheil, 1915, l n t . ent. Z., 9 : 89, 101, Test-fig. 3. [Holotype d. Algeria,

Constantine dep., Sidi Okba, 4 . v i i . 1917, (Kheil). Sational Ifuseum, F'rague.] Syn. n.

DIAGSOSIS 9. 1. Aedeagal valves slender, upturned, not decurved posteriorly (Text-fig. 13 : B. p. 311). Profile when covered by ectophallic pocket as in Test-fig. 3 : S. p. 296. Cingular

c

3

I)

1 L

A R E \ - I S I O S O F T H E G E S C S C A L L í P T d i l l U S S E R V I L L E 32'

valve (Test-fig. 13 : C, a. p. 311) and lateral accessory processes of penis valves (Test-fig. 13 : C, 1, and B, 1. p. 31 1) pointed and membranous, latter curling dorsally to form a tube involving cingular and penis \dves (Test-fig. 10 : E, 1 and m). Postero-dorsal part of cingulum and rami of cingulum (Test-fig. 13 : B, g. p. 31 1) rugose and swollen, though not overhanging tip of penk (as in C . coeZesFviensis, p. 343). Dorsal ectophallic membrane plate with simple laminar posterior median edge (Test-fig. 13 ; E. p. 31 1), being only slightly thickened and rugosely sculptured in this region ; plate almost as wide as long (cf. Text-fig. 13 : F. p. 311). Rami of cingulum, viewed from a posterior aspect, converging upwards (Test-fig. 13 : B. p. 31 1).

2. Cercus variable apically (as in Test-fig. 2j : A-S). Xlways three apical lobes, though median lobe in some Korth African material may be ven; weak. 3. Tegmina often reduced in size ; Rs reduced to two branches in European forms, usually

three branched in Sorth African series. 4. Welanic forms (Sorth African), or near melanic forms (E. Pyrenees) occur. Othenvise

brownish grey with darker mottled spotting. 5 . Inner side of posterior femora typically with inner median area pink, this colour extending

on to loiver inner carina (in C . titvanictts never doing so). Three inner femoral spots (usually separate), median one largest, al1 extending to some degree below inner dorsal carina (Text-fig. 2 0 : H and E;. p. 324) (in C. ttcrarricrrs never doing so, Text-fig. 2 0 : J), In darkly pigmented material from S. Xfrica (Atlas), anterior and median spots may fuse, median spot in N. African material being larger in proportion than in European material even if unfused.

6. Posterior tibiae variable in colour, from pale flesh pink to crimson. Often paler externally than dorsally or internally, but can be uniformly pigmented. May be suffused with black giving dark mauve appearance (e.g. in menees and N. Africa).

7. Hind wings pale pink to deep crimson, pigmentation covering basa1 half to three-quarters of wing or more. Anal fan always more deeply pigmented than rest of wing. Folded wings usually not surpassing apices of folded posterior femora.

8. Tegmina coarsely spotted, though contrast not always marked, spots being merged into a rather uniform brown except at apes. In dark specimens with contrasting colouring, spots may become seriallp spaced blotches running diagonally across tegmen.

9. Diagnosis as for male. Bulky insects with tegmina which taper conspicuously in their apical 2/3, and whose apices just surpass or just fa11 short of knees of folded posterior femora (see key to 99, p. 310).

TABLE 1.

~ Tegrninal opices surpassing(L),level with (=),

FIG. 18. Trgminal-femoral relationships for Sorth African material of C. wuttenzylianus (Pantel).

,

322 K. D . J .4GO

I

I '!

I 5mm. I X 9 A

2 111111. ,

,

323 A R E \ - I S I O S OF THE G E S C S CALLIPTddICS S E R V I L L E

NEAS~REJIESTS Males Fe m a 1 e s A r \ I h

-l

S O . Range JIean so. Range Mean Head width . j Y 3 3 8-4.7 4 ’ 1 j . 118 j ’0 -7 .1 j.7S Femur length . -- / I 10.s-1j.S 11.80 . 116 14.j-2j.0 29-03

10.2-1s.j 14.80 . 1 1 7 11.8-31.j 21.56 Tegminal length . -- ,/

Total length . -- / I 17’3-2S.1 21-43 . 117 21.6-41’9 31.j1

DISTRIBUTIOS. Lying in two series of populations, separated by the Straits of Gibraltar (see Text-fig. 6, p. 301), this species penetrates the High and Riiddle Atlas mountains of Jlorocco, the uplands of Algeria, Tunisia, and eastwards to northern Libya, while in Europe it occupies southern and Eastem Spain, southem Mediter- ranean France, and lower valleys of the Maritime -41~s.

FRAXCE : Pyrénées-Orientales, 3 0,4 9, viii-xi ; Hérault, I $, vii : Var, 11 3, 26 Q, viii-ix. SPAIX : Cadiz, 2 9, vi; Gerona, I $!)--. MoRocco : Volubilis, I 9, vii ; Moyen Atlas, up to 1,700 m., 3 3, 5 9, vi-ix ; Hauts. Atlas, 1,300-1,900 m., 21 3, 26 9, vii-viii ; nr. Tangier, 2 3, 5 9, viii ; nr. Fks, I 9, viii. ALGERIA : Oran, g 3, 6 9, vi ; Hauts Plateaux, 1,100 m. I 3, -5 9, vi-ix ; Chardia, 2 6, 8 Q, s ; Alger, 7 3, g Q, vii-jx ; Constantine, I d, 2 Q, vii. TUNISIA : Gafsa, I d, iii. LIBYA : Cyrenaica, 4 d, 11 9, vii-ix.

DISCL-SSIOX. This species shuns either high altitude and cold humid conditions, or extreme dq-ness. Consequently although often sympatric with C. barbarus, C. italicus, and C. stdalpiiztrs, it has a more restncted distribution than the first two, and does not enter montane habitats with the last. Usually in a minority,it has been found in swarm proportions, forming up to 90% of the Calliptamus populations present. Typical localities are found on old stabilized dunes, or among untended vineyards.

C. okbaemis ic synonymized in this paper with C. wattenwylialzzts since in the morphology of the male genitalia both are identical. Both species show :

(i) tendency to formation of melanic forms, (ii) shortening of tegmina with altitude, (iii) same variations of epiphallus, femoral markings, and tegminal renation, (iv) same range of size as measured by total length and head width.

P~L~TERIAL EXAJIIXED.

E:

t

c

FIG. rg. A. Lateral aspect of entire male of C. cyenaictrs sp. n. to show esternal tasono- mic characters used in this genus. 1. Tegmen or foreiving ; 11. Swollen 9th and 10th abdominal tergites : 111. Supra-anal plate ; IV. Cercus-used as a clasping organ during copulation : Y. Ectophallic pocket covering, penis valves (cf. Fig. z : X) ; VI. Sub- genital plate ; 1-11, O ‘ Knee ” of posterior femur : VIII. Posterior tibia and spines ; IX. Posterior tibia1 spurs (outer pair). E J . Dorsal aspect of pronotum in a series of specimens of C. cyeiznicus sp. n. (from Libya, Cyrenaica prov., Shahhat), to show range of colour polyniorphism. B. var. mn~,gi~dlzts in its most striking form. C. paler form of B., witli central area ginger brown. D-F. commonest forms in any population. These predominate almost completely in boreal populations of C. itnliczcs (L.) and C. I>oyDnrzis (Costa). G-J. uniformly pigxnented forms, G. having a disc to posterior transverse sulcus slightly darker in colour. G. dark and light red-brown, J . pale buff, H. intermediate between these two estrenies.

b i

324 ii. D . J A C O

brown and dark brown

A 0.

dull dark brown ochre

white -

J K

FIG. 20. A-C. Lateral aspect of head in 3 individuals of C . cyrenaicrts sp. n. to shoLv coiour polymorphism. (N.B. 5 mm. scale refers to diagrams A-D). D. Lateral aspect of pronotum in some species, to show general pattern typical of genus as a whole. E-G. Posterior tibia1 spurs in the genera Calliptamrrs Serville (E), Caloptenopsis 1. Bolivar (F) (cf. Fig. 3 : U), and .4corypha Krauss [G). H-K. Inner surface of left posterior femur. J. pale male form in C . tirranicrts Tarbinsky. E;. dark female form in C. wattenzqlianris (Pantel) from south-eastern France. H. intermediate form with medium development of melanic areas in female of C . ziaftentiylianzis (Pantel) from eastem French Pyrenees. (colour reprecented as in Fig. g : P-L).

i

334 S . D . J . \GO

>fEASUREMESfC Males Females 7- - S O . Range 3Iean K O . Range hfean

Head width . . 497 2. a-4.8 3'69 . 897 4.2-6.8 5.26

Tegminal length . 495 9.3-23'6 14.46 . 896 10.3-34.8 23-24 Totallength . . 494 10-5-30.5 20.10 . 855 18.2-46-j 31.66

Femur length. . 492 7'8-17'4 IO.8j . sj j 12'0-22'7 14'4j

DICTRIBCTIOS. As indicated for species, excluding area occupied by subsp. pnZnesti.ite?tsis Rme. (see Text-fig. g, p. 305.). The range of the nominate siibspecies into the Canary islands mentioned by \Yillemse (1936) was based on information supplied by Ramme, who said he had collected material from there. As far as is known C. Plebeiirs is however the only species found on those islands.

3L4TERIrlL EXAYISED. PORTUGAL : Alto Alentejo, g Q, - ; Trans-os-Montes e Alto Douro, 5 $2, Viii-xi. SPAIK : Segovia, 4 8, 6 $2, viii ; Murcia, 4 $, 3 Q, ix ; Granada, 1,800 m., I $2, ix ; Aviia, I d, 4 $2, v ; Alicante, 2 $, 2 $2, ix ; Santander, I 8, 2 $2, ix ; Gerona, 4 8, 2 $2, ix ; Barcelona, I 8; 5 $2. viii ; Mallorca I., I 8, 7 $2, xi. FRASCE : Pyrénées-Orientales, 15 8, 15 $2, viii-ix ; .4ude, 13 $, 4 $2, viii ; Puy-de-Dome, 19 8, vii ; Lozere, I 8, vii ; Hérault, 17 3, 10 $2, vii-viii ; Bouches du Rhone, 2 b, I $2, viii ; Var, 16 $; 17 $2, viii-x ; Aipes-3íaritimes, 35 $, 52 $2, vii-ix. ITALT : Lecce, 21 $, 12 $2, ix-xi ; Foggia, 2 5, I $2, vii ; Imperia, I 8, 2 $2, \1i-x ; Potenza, 2 d, I Q, viii ; Grosseto, Go m., I 8, 3 $2, ix ; Napoli, 7 d, 7 $2, vii-ix : Basilicata, 420 m., 5 $, 6 $2, ix ; Sardinia I., 2 8, 2 $2, vii ; Lipari I., 3 8, 7 $2, viii ; 1' di Tremiti, S. Nicola, S. Domino, Pianoso, Capraia, 4 8, 4 9, vi-ix ; Roma, I d, I $2, - ; Ban, 2 8, 2 $2, ix ; Sicily, 2 d, viii. JUGOSLAVIA : Srbija, g $, 21 $2, viii-x ; Vojvodina, 60 m., 2 $2, viii ; Makedonija, 100-200 m., I 8, 2 $2, viii. Hurl- GARY : btwn. Tiza and Dunar R., 5 d, 5 $2, vii. GREECE : Rlakedhonia, 2 8, 10 $2, vi-viii ; Thessalia, 2,000-2,300 m., 2 8, vii ; Stereá Ellás, 13 $, 23 $2, \di-viii ; Sporadhes I", 400 m., 10 Q, vii ; Rodhos I., 2 d,2 $2, viii ; Pelopónnisos, g00-1,ooo m., 24 d, 27 $, vii ; Kriti I., up to 1,700 m., 12 0, 27 $2, vii-viii. CYPRUC : T'ery long series from ali parts. hL4LT.4 : I 9, -. JfoRocco : 'rioyen -Atlas, up to 2,5jo m., 7 5, 27 Q, vi-is ; Hauts Atlas, up to 1,700 m., 3 3, 4 9 , vi-ix : Tangier, I 9, viii ; Rabat, 6 8, 4 1, vi ; Port Lyautey, 2 5, 2 9, viii ; Ceuta, I j, 2 $2, - ; Ifrane, I $2, wii. -\LGERIA : Alger, up to 1,300 m., 8 8, 2 j $2, vii-ix ; Chardaia, 46 d, 92 $2, x ; Oran, 2 $2, vi ; Touggourt, 5 $2, vi ; Constantine, I 8, I 9, vi. TUNICIA : Gabks, I 3, I $2, vi. LIBYA : Tripolitania, up to So0 m., 7 $ , S $2, v-xi. TURKES : Ankara, 4 $, 3 Q, viii ; Tekirdag, I $, 2 $2, viii ; Amasya, I 3, viii ; Rize, 3,200 m., I $2, viii ; Urfa, 11 $, 4;1 $2, vii ; Hatay, 3 5, 23 9, vii ; Makkiri, 1,600 m., 3 $2, viii ; Seyhan, 1,500-1,900 m., 2 $2, ix ; Gasiantep, I Q, - ; Istanbul, 2 3, 6 $2, vii-x ; Zouguldak, 10 5, 2 $2, viii ; Jíugla, 2 $, I 9, vii : Nanisa, up to goo m., 11 d, 11 $2, viii ; izmir, 7 8, 6 $2, vii-viii ; Denizli, i,jOo-i,gOo m., 2 j, 9 9, viii. SYRIA Latakia, 13 d, 6 $2, c-iü ; Damascus, I d, vii ; Tripoli-Homs, 3 9, vi. LEBAKON : Hasbaya, I Q, vi : E. of Beirüt, 2 5, I 9, vii : -Amyün, I Q, Tii. ISRAEL : Samaria, I 2, si ; Haifa, I $, 2 9, xi ; Northern distr., 2 3, j 9, vii-is ; Plain of Sharo, 5 $2, i s ; Southem distr., Negev, I 5, I 9, vii-viii ; Southern distr., Beersheba, j j, 3 $2, v-vii. JORDAS : E. Ramallah, 2 9, vi ; El Ghor, Damiya, G z, 6 T, vi ; Irbid-3lafraq

...

...

A R E V I S I O S O F T H E GEXUS C A L L I P T A M U S S E R V I L L E 333

rd., 6 $, vi ; S. of Mafraq, I 5, vi ; IY. of El Salt, 3 8, 5 9, vi ; \Ya& Shueib, 2 3, vi ; nr. Nadaba, I 5, viii. EGSPT : Lower E a p t , I O, - ; Lower Egypt, Faiyíim, I d, vii : -, 2 9, vii-viii. IRIQ : lIesopotamia, Baghdad, 5 3, 3 9, vi-sii ; Mesopo- tamia, liendali, I 6, vii ; Iíurdistan, I o", v. IRAS : JIarkazi, up to 2,100 m., 2 $; 7 Q; 1%-si ; Hashtom, up to 2,400 m., I 8, 2 $2, v-x ; Sheshom, j 8, 10 Q , vii-ix ; Haftom, up to 1,800 m., 3 3, 5 $2, v-vi ; Chahirom, up to 1,700 m., I $, 2 $2, viii-x , Dovvom, I $2, vii. AFGHAXISTAX : Kabul, go0-2,500 m., S 6, 49 9, vi-s ; Badakh- shan, I,SOO-~,~OO m., 14 3, Sg 9, vii-viii ; Eactem prov., Kuristan, 1,100-2,700 m., 6 3, 16 9, vii. IV. PAKISTAX : Jammu and Kashmir, up to 2,600 m., 5 $, 7 9, ix-x ; Chitral, 4 3, S 9, v-is ; Baluchistan, I 3, 2 $2, vi-vii. BULGARIA : Sv. Vlas, I 3, I Q, viii. U.S.S.R.: Ukraina S.S.R., 2 3, I $2, vii; Kazakhctan S.S.R., 30 $, 34Q, v-ix ; Daghestan A.S.S.R., I 8, 4 9, vii-viii ; Azerbaydzhan S.S.R., I $2, vii ; Tadzhikictan S.S.R., up to 2 ,O jO m., I $, I $2, vii-viii ; Turkmeniya S.S.R., I 8, 4 9, vi ; Armeniya S.S.R., 7 8, 42 $2, vii-ix.

Calliptamus barbarus palaestinensis Ramme stat. n. Calliptaílttts palaestinensis Ramme, 30.vi. 1930, i2Iitt. zool. Xits. Berl., 16 : 395 ; Bodenheimer,

vii. 1930, Monogr. angew. Ent. 10 : 62. [Holotype d. Israel, Palestine reg., Ben Shemen, 1925, (F. Bodenheimer) Berlin Mus.]

Calliptamus palaestinensis evythrocnemis Ramme, 1951, Mitt. 2001. Alirs. Berl. 27 : 313. [Holo- type d, Syria, Beirüt, Xahr el Kelb, iii. 1925 (2iIiillev). Berlin bIus.1 Syn. n.

DIAGNOSIS. See key, p. 332.

WEASUREJIENTS Males Females - - Head width . . 24 3.4-4.5 3'71 . 53 4.6-6.4 5'51

?FO. Range Mean S O . Range Mean

Femur length . . 24 9'4-14.2 10.82 . 52 1j*0-22.0 18.05 Tegminallength . 21 9.2-14.0 11.72 . 53 15'4-27'5 20'93 Total length . . q 14.2-26.2 17.17 . 53 23.7-35.7 30.00

DISTRIBUTIOS. See key, p. 332. MATERIAL EXAJIISED. LEBANON : Laqlaq, 1,500 m., 11 8, g 4, vii ; Bruaman,

2 9, 2 Q, vii ; (Intermediate forms) Beirüt, 2 3, 6 $2, v-xii ; Baalbek, z Q, vii. ISRAEL: GaIiIee, I$, xi ; Korthern distr., 7 $, 5 Q, s-xi; Haifa, I $, 19, ?u; Jerusalem, 6 3, 20 9, vi-si. JORDAX : El Ghor, Jericho, I 0, s.

>

Calliptamus cyrenaicus sp. n. DIAGXOSIS d. I. Cingular valve elongate, membranous (Test-fig. 25 : G. p. 339), reaching

well beyond tips of lateral accessory processes. Tip of aedeagus, as seen from above, narrotver when compared with width of cingular arch than in C . barbarus, i.e. Cingular arch width/width of aedeagal valves, for C . cyvennicrts 1*97-2*20, mean 1-54 ; for C . barbartts 1-20-1.73, mean 1-46.

1

2 . Lower apical cercus lobes tend to be reduced (Test-fig. 24 : 1, 11, VI. p. 336).

336 h’. D. J . 4 G O

l 7 mm.

A R E V I S I O S O F T H E G E N U S CALLZPTAdlUS S E R V I L L E 337

3. Tegminal apices never surpassing knees of folded posterior femora. Distinctly tapered apically (Text-fig. 19 : X. p. 322). 4. Inner femoral spots separate, median spot rather large, square, nearly reaching lower inner

canna. Posterior tibiae and inner side of posterior femora deep ruby to pale scarlet. 5. Anal fan pale to deep scarlet. Area anterior to this hyaline, colourless, with strongly

pigmented brown veins. Kell developed colour polymorphism (Test-fig. 19 : -4-J. p. 322, and Text-fig. 20 : A-C. p. 324).

9. Unlike sympatric C. bavbarits never have solid black femoral spot. Sympatric C. wntletr- wylinnus females differentiated on larger size, their measurements being as follows :

so. Range hIean Head width : . . 15 5 * 7-6 ‘ 4 6.11 Femur length . . 15 19-5-22.5 20.73 Tegminal length . 15 19-8-25.7 22.79 Total length . . 15 30.6-37-0 33-41

MEASUREMENTS Males Females L - r \

Head width . - 9 3.6-4.1 3.83 * 21 5.1-5.9 5’44

NO. Range Mean NO. Range Mean

Femur length . - 9 9’4-11’5 10.75 . 22 15.8-19-2 17.73 Tegminallength . g 9-9-12.0 11.07 . 21 15-0-20.8 18.32 Total length . . g 15.1-18.1 16.91 . 21 23.2-30.2 27-19

DISTRIBUTION. MATERIAL EXAMINED. Holotype g, Libya, Cyrenaica prov., Slonta, Jebel Akhdar,

r.vii. 1953 (K. M. Guichard), Brit. hluc. (nat. Hict.). Paratypes, 2 8, 7 9, came data and depository. LIBYA : Cyrenaica prov.; Slonta, Jebel Akhdar, 3 8, 7 9, I . vii ; Shahhat, 500 ni.,

Only in Northem Libya (Cyrenaica) .

i

.-

FIG. 24. Outer apical surface of left male cercus. [u) 1. C. cyvenaictis sp. n. Libya. Cyrenaica prov., Tokra. (b) 11, 1-111, XI, XIV. C. barbavzis barbnrzrs (Costa) showing trends as those seen in C. stibulpinirs sp. n. (XVII). 11. hlorocco, Atlas mts., Ait Bou Guemmez, 6,000 f t . 1-111. Spain, Segovia prov., Sierra da Guadarrama. XI. Spain, Santander prov., Picos de Europa. SIV. France, Puy de Dome, 16 kms. S. of Le Puy. (c) 111-VII, IX., XII, XIII , XV, XVI. C. barbavzis barbnrzis (Costa). 111. Morocco Moyen Atlas, Aguelman Sidi Ali Ou hlahommed. 6,500 ft . IV. JIorocco, Moyen Atlas, Col du Zad, 2,000 m. 1-. JIorocco, Rabat. VI. Algeria, Alger dep., Chrea, nr. Blida, 1,30Ó rn. VII. Algeria, Ghardaia dep., Djelfa. IX. Spain, Segovia prov., San Rafael. SII. France, F’yrénées-Orientales, Banyuls-sur-Mer. XIII. France, Pyrénées-Orientales, Vernets-les-Bains. X\-. France. Iiérault, Palavas-les-Flotc. S V I . France, Provence prov., 1‘ar dep., Fréjus. (d) XYII. C. subalpimrs sp. n. France, Alpes-Naritimes. (e) XIX. C. plebeizrs PValker), Canary Islands, Gran Canaria 1. (f) X-S. C. wattenwylianzrs (Pantel). France : A. 1-m. St. RIaxirnin distr. B. Var, Wounes-les-Jíontrieus. C. Hér- ault, Montpellier. D. Pj-rénées-Orientales, Banyuh-sur-Jíer. E. Pyrénées-Orientales, Yernets-les-Bains. Cpaiii : F. Gerona prov., Port Bou, Cataluna. Libya : G. El Xarj and Slonta. JIorocco : H. Tangier and Ras el RIuar. J . Rabat. K. RIoyen Atlas, Tirn- hadit, j,OOO ft., L. Jíoyen Atlas, Timhadit and RIouldirt, JI. Atlas SIts., Ait Bou Guernmez, 6,000 f t . S. Nouldirt (and Algeria : Volgrove). O and P. Atlas Blts., Ait Bou Guemmez, 6,000 fr. -4lgeria : (2. Oran dep., Mascara. R. Oran dep., hlascara ; and Alger dep., Boghari. C. Alger dep., Boghari. (g) XL’III. C. siciliae Ramme.

.

338 S . D . J A C O

I 8, 22.x. and 28 .ix ; Shahhat to Tokra, 3 $, I.X ; Derna, zoo m., z 9, 3 ? , 2 3 .ix ; Saff Safi, z 9, 26.ix ; Latrun, 10 km. E. of Ras Hilal, I 0, S.iv. ( A . F. Sladden, 1. A . D. Robert.co?t, K . JI. Gziichnrd). +U in Brit. Ulus. (Nat. Hist.).

DISCCSSIO?;. Probably evohed from C. barbarus (see diccussion, p. 292).

Calliptamus subalpinus sp. n. Cnlliptniizzrs icfeericzrs Serville, auctt. (misidentiíícation).

DIAGSOSIS 8 . I. Rather short, weak, apically outcurved penis valves (Text-fig. 25: E. p. 339). Lateral accessory processec of penis valves similar to C. barbariis (Text-fig. 10 : E. 1). Orientation of aedeagus as in latter species (Test-fig. 3 : T. p. 296). Cingular valve with its apes leve1 with a line drawn to touch posterior apices of lateral accessory processes of penis valves.

2. Cercus apes variable but ranging from that shown in Test-fig. 24 : XVII. p. 336, to that of C. siciliae (Text-fig. 24 : XVIII). Usually near second type. 3. Tips of folded tegmina never surpassing knees of folded posterior femora. Tegmina tapered

in apicai 213. 4. Darkly coloured ; pronotum type E (Text-fig. 19. p. 322). Little colour polymorphism. 5. Posterior femora with three separate black or dark brown inner spots, middle spot usuaiiy

largest. Often paralleling C. italiczis very closely a t high altitude. Spots may be faintly pigmented, with little black pigment below upper inner canna. 6. Posterior tibiae dull ruby-red to red ; this colour also on lower inner femoral carinae. 7. Hind wings colourless or flushed with pink basally (polymorphs distributed a t random

geographically) . 9. For separation from other syrnpatric Cnlliptamzis spp. see pp. 318 and 328 ; also key p. 310.

C. uattenwylianzcs never occurs above joo metres, and never has colourless hind wings. A larger insect [compare measurements, p. 323 with those below). C. subalflinzis and C . siciliae females inseparable except on distribution.

>IEASCREMENTS Males Females h L

f f \

S O . Range AIean XO . Range Blean

Femur length . . Sj 7.6-10.5 9-12 . 147 13.4-16.9 15.37 Tegminallength . Sj 6.4-11.3 8.77 . 147 13.7-19*9 16-80 Totallength . . 85 10-8-16.9 13.72 . 147 20-tL28.6 24-98

Head width . . Sj 3.1-3.7 3'34 147 4.13'3 4-71

DICTRIBUTION. Present in the Waritime Alps and south-eastem France (up to 1,200 metres), it is unable to penetrate the southem Italian Alps, but occupies the higher Ligurian Alps and the Italian peninsula south of the Po vaiiey. (Test-fig. 6,

h i . 4 ~ ~ ~ 1 . 4 4 ~ EXAJIISED. Holotype 3, FRAKCE, Alpes 'Iíaritimes, Valdeblor St. Roch,

Pnuntjpes, 23 8, 10 0, carne data and depository. FRASCE : Alpes Naritimec ; Biot, 200 m., I j, z 9, viii ; Le Rouret,,300 m., 20 d,

19 9, IZ .ix ; Sarre valley, Goo m., 19 6, 39 9, ix ; Ct. llartin-Vésubie, 21 6, 34 9, ix ; St. Dalmas de Tende, 1,100 m., g 8, g 9, 5 .viii : i'aldeblor St. Roch, 1,120 m., 24 d, 11 9 , s . viii ; Thorenc, 7 d, 20 9, i s ; Le Souquet, 2 .ix - Var ; Bagnolc, 19 8, 16 9, x ; RIéounes-les-Nontrieus, K. of Toulon, r 8, 2 9, 1 . i ~ : Blontouroux, 350 m., 4 d,

P. 301).

1,120 m., 5 . ~ 3 . igj8, ( N . D. Jago). Brit. RIus. (nat. Hist.)

339 A R E V I S I O S O F T H E G E S C ’ S CALLZPTAdfUS S E R V I L L E

b’ 0.5 rnrn.

FIG. 25. Apical part of phallic complex in males of “ barbnrzrs ” complex. Al1 diagrams show dorsal aspect, except F. which showvs view from right side. (u) A-D, F, and H. C. burbnrris bnvbnrics (Costa) : A. Topotype, Italy, nr. Napoli, Ponte de Mare. B. Spain, Segovia prov., Sierra de Guadarrama. C. Jordan, W. of El Salt, IChor Kabid, D. Jugo- slavia, Srbija reg., Deliblatski san&, Pannonia. F. Jordan, W. of El Salt, Khor Kabid. H. West Palristan, Jammu and Iíashmir state. Srinagar, 3,500 f t . (b) E. C. strbulpinzrs sp. n. France, .Upes-AIaritimes, Valdeblor. (c) G. (and inset showing cingular pouches) -C. cyremicits sp. n., Libya. (d) J. C. bnrbnvtcs pnlaesiinensis Ramme, Lebanon. (e) K. C. siciliae Ramme, Sicily.

340 N . D. J A G O

23 9, x : ( N . D. Jago, 31. Korsakoff, D. R. Rngge, B. P. Uztarov). Brit. 3111s. (Xat. Hist.). ITALY : Genova, Chippiato, 6 3, 3 $2, 6.x and 11 . i s ; Lipria, Chiavari, I 8, I $2, ix ; Toscana, Stiava, nr. Yiareggio, 1-40 m., I d, 3 9, 16.ix ; Potenza, i'arco di Pietrastretto, nr. Potenza, 850 m., I 9, 3 9, 2 G . k ; Basilicata, Venosa, 420 m., I 9, 30. i s ; Cosenza, Calabria, I 3, x ; Cosenza, Cozzadi JIantielia, I 8, 3 9, 3r.viii ; Roma, st. I'ito Zanon, I d, I 9, -; Puglie, Maglie, 70 m., 30 3, 29.ix : Salentina pen., Taranto, Martina Franca, I 3, g.ix ; Bari, Murgia, Serraficaia mt., I 9, n . i x ; Lucania, Ateiia, 2 8, 12. vii ; Bari, Aitamura, Xurgia Parisi, I $2, 22.vi . (F. Capra, &l. Su@, F. E. Zeztlter). Representatives in Brit. JXus. (Nat. Hist.).

DISCUSCION. This speciec cannot take the name C. ictericw (for discussion see p. 332). Although very close to C. siciliae, the species differs in the form of the penis valvec. The species is probably derived from C . barbarus, many of whose upland populations in Spain and France show similar externa1 facies.

Cafliptamus sicifiae Ramme, stat. n. Calliptamus abbreviatus siciliae Ramme, 1927, Eos, 3 : 166. [Lectotype d, Sicilia, Messina

prov., nr. Messina, Colle S. Rizzo, 2-400 m., 27.vii.1924 (Ramme & Richter) Berlin Mus. Only types studied.]

DIAGNOSIS d. I. Penis valves with widest part exposed beyond posterior edge of cingular

9. Separable only on distribution. Otherwise like C. szibalpinus. valve (cf. Text-fig. 25 : K. p. 339). Other characters as for C . szrbalpznus (diagnosis, p. 338).

MEASUREXESTS Males Females - f No. Mean &-O. Mean

Head width . . 2 3.3 and 3.6 3-45 . 2 4.gand 5 . 2 5 'O5 Femurlength . 2 9.2 and 10.1 9.65 . 2 16.5 and 17.8 17.15 Tegminallength . 2 8 .oand 8.6 8.30 . 2 16.1 and18.1 17.10 Total length . . 2 12.8and 14.2 13.50 . 2 24.7andz7.8 26.25

A \

DISCUSCIOX. A lectotype has been chosen from Ramme's syntypes, and is now returned to Berlin Museum. The small amount of material may indeed mean that these genitalia are atypical, in which case C. stcbalpilzzu must be sjnonymized with C. siciliae.

Calliptamus tenuicercis Tarbinsky, 1930 Calliptamzis tenziicercis Tarbinsky, 1930, Bzill. Acnd. Sci. U.R.S.S. : 180. [Holotype $., Iran

Chaharom prov., Gandzha, 2 .viii. 1928, (Tavbinsky). Leningrad h.Ius.1 Cnlliptatnris iraniciis Ramme, 1930, d l i t f . zool. Jl i i s . Berl., 16 : 395. [Syntypes 22 d, 27 0 ,

Azerbaidjan, Ordubad, 3 .viii. 1927 ; Armenia prov., Ecmiadzin, ?o.viii. 1927 ; Magos, 12-800 m., Zj.\'iii. 1927. Berlin hlus.]

Calliptamiis iranicits auruntincus Ramme, 1930, Ibid. [Unspecified number of syntypes, Tiflis to Mzchet. Berlin Mus.] Syn. n.

Callipfnmris persa Gvarov, 1938, Ann. ilPag. nnt. Hist., ( I I ) , 1 : 371. [Holotype d, Iran, Seshom prov., Rlasjed-Sol5yman, nr. Ahvaz, vi. 1937, (S.\'.P. Pill). Brit. Mus. (nnt. Hist.).] Syn. n.

'

A R E V I S I O N O F T H E G E N U S C A L L I P T A M U S S E R V I L L E 34'

Calliptamzcs ienuicercis iraczis Maian, Igji, Sbor. eñt. Odd. nár. Mus. Praze. 27 : 384. [Holotype $, Iraq, hfesopotamia reg., Baghdad, (Kalalooa). Sational bIus., Prague.] Syn. n.

Calliptamais tentriccrcis syriacus Ramme, 1951, ilfitt. 2001. Mzis. Berl., 27 : 310. [Syntypes I o", 3 $2, Syria, (Ehrenberg). Berlin hius.] Syn. n. DIAGSOSIC $. I. Weak pointed penis valves (Text-fig. 15 : E and F. p. 31 j), almost verticaliy

orientated. nearly hidden by cingular valves from above (Test-fig. I j : E. a). Lateral accessory expansions of penis valves auricular, hiding al1 but tips of penis valves when viewed laterally (Text-fig. 15 : D, m. p. 315). Cingular valve thick and flattened apicaily (Text-fig. 15 : F, a. p. 315). Dorsal ectophallic plate broader than iong ; posterior median lobe laminar and uptumed. C. coelesyriensis differentiated by shape of cingular arch, cingular valve, and dorsal ectophallic membrane (Test-fig. 16 : C, c. p. 317). 2. Cercus (Test-fig. 17 : H, J. p. 319) with apical lobes usually well differentiated. Ratio of

length to maximum depth near apex, 4.0 plus (like C . barbarus, but see C . balucha, p. 342). 3. Tegmina with apices leve1 with, or falling short of, (occasionally surpassing) knees of folded

femora. Rs with up to 4 branches. 4. Posterior femora yellow on inner side ; often dull pale ruby between inner carinae. Inner

femoral spots separate to fused, if completely so then no orange and grey-ruby pigment just posterior to black area (cf. C. barbartrs p. 314 ; see Text-fig. 22 : B. p. 330).

5. Posterior tibiae yellow, pale dull orange or orange. 6. Hind wings ruby in basa1 4 to +-rest colourless with darker veins. 7. Tegminal colouring as for C . barbarus ; '' marginellus " forms common. $2. For separation from C . barbarus see pp. 314, 328 and 331.

i M E ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ Males Females A A

I I \ h-0. Range Mean No. Range Mean

Femur length . 99 8.2-15.6 11-75 . IIO 12*0-22-0 17.02 Tegminallength . gg 9.0-20.7 14.59 . 111 13-638.7 21.40 Total length . . gg 13.5-28.2 20.39 . 111 20.2-38-8 30'23

See Text-figs. 8 and 9, pp. 304 and 305 respectively. The species inhabits an area where mean annual rainfali is generally less than 25 in. It thus penetrates semi-desert environments more deeply than does C. barbarzu which is lesc xerophilous.

~ ~ A T E R I A L Ex4hfINED. TURKEY : Ankara, 5 8, 8 $2, viii ; Antalya, 5 8, 6 9, - ; Istanbul, I 3, I $2. viii : Seyhan, I 9, - : Urfa, 4 3, 27 $2, vii ; Nigde, 1,300 m., 2 $2, ix ; ali foliowing va^. aurafitipes Mersin, 1,goo m., 10 9, viii : Denizli, 1,280 m., I $, viii ; Hatay, I $2, yiii. LEBANON : Bekaa Bega, I $, vii. ISRAEL : Northern distr., 3 8, 2 $2, i v ; E. of Nablus, I d, v ; Southem distr., Negev, 4 8, 2 9, vii. JORDAN : Ain el Basha, I 8, vi ; El Ghori 16 8 10 9, vi : Irbid-Mafraq rd., I $. I $2, vi : nr. El Salt, g 8, 8 9, vi; Wadi Shueib, 10 8, 6 $2, vi. IRAQ: Mesopo- tamia, Baghdad, I 8, I $2, vi. IRAN : Chahárom, 2 3, 2 $2, viii-ix ; Marhazi, I 8,5 Q, vi-ix ; Dovvom, I 8, vi ; Hashtom, 8 3, 6 9, iii-x ; Haftom, up to 2,500 m., I d, ix ; Seshom, 14 8, g 9, vi. AFGHANICTAS : Kabul, 2,400 m., 2 8, I 9, vii. U.S.S.R.: Armeniya S.S.R., 2 8, 2 9, vii-ix ; Azerbaydzhan S.S.R., I 8, I 9, vii ; Daghestan A.S.S.R., 2 8, 5 9, viii ; Gruzija S.S.R., I d, I $2, v.

DICCUSSIOS. Phallic morphology is very unifonn throughout the range of distribution of this species, colour variation being explicable on the same arguments as those used for C. bnrbnrzis (pp. 301-306). The new synonjmc are based on material

Headwidth . - 99 3 * 2-4 *8 3-92 . 115 3'94.2 5-17

DISTRIBUTIOS.

342 S . D. J A C O

which ali belongs to the orange-legged populationc in the east of the range (see Text-fig. 22 : B, p. 330). The species ceems most closely related to C. balzrcha and C. coelesyrieiasis.

Calliptamus balucha Uvarov, 1938 Culli~5iamzrs bulucJzu Uvarov, 1938, Ann. J fug . nat. Hist. ( I 1), 1 : 371. [HolotFpe $, West Pakis-

tan, Baluchistan reg., Ziarat, 8,000 ft., 6.viii. 1929. (J. W. Evans) Brit. Mus. (Nat. Hist.).]

DIAGNOSIS d. 1. Penis valves of lowland forms like C . tenzticercis (Text-fig. 15 : G-J), but in montane forms (above 2,000 m., e.g. Chitral and K.W.F. prov., LV. Pakistan) more like C . barbarzcs (Text-fig. 15 : A-C. p. 31 5). Epiphallus ratio of width across loiver corners to depth a t centre never more than 2.0 (C. tenuicercis, ratio 2'1-2.9). Epiphallus heavily sclerotized (Text-fig. 15 :

2. Cercus (Text-fig. 17 : H, J. p. 319) with ratio of length to maximum depth at distai end

3. Tegminal apices never surpassing knees of folded posterior femora. Often so reduced that

4. Inner posterior femoral colour dull greyish ruby to pale ruby-red. Black femoral spots

5. Posterior tibiae dull greyish ruby to pale ruby-red.

NI.

never more than 3.8 (range 3'30-3'60), cf. c. tenuicerck, p. 341.

apices do not reach middle of folded posterior femora (extreme montane forms).

separate ; median largest and often diñuse.

KEY TO SUBSPECIES

I. Penis valves (Test-fig. I j : J, m. p. 31 j) relatively weak. Cingular valve like C . tenuicercis (cf. Test-fig. 15 : H, a and F, a. p. 31 j) its width apically being well over half width across outer edges of lateral processes of penis valves. Penis valves half hidden by lateral accessory processes from side view (Text-fig. I j : J. p. 31 5). Tegminal length to femoral length ratio. ^o-96 8, never less than 1.0 in 9

balucha balucha Uvarov (p. 342) -. Penis valves (Text-fig. I j : A-C, m) relatively strongly developed, reminiscent of C.

barbuvus (Test-fig. zj : F. p. 339). Cingular valve only just wider apically than half width across outer edges of lateral accessory processes (Text-fig. I j : A, 1) . Penis valves clearly visible in lateral view, hardly hidden by lateral accessory pro- ceses (Text-fig. 15 : C). Penis valves with flattened and moderately outwardly expanded upper edges (Test-fig. 15 : B, m). Tegminal length ratio 0'62-0'92 8, 0'66-0'96 9 . . balucha brachypterus pirsh) (p. 343)

Calliptamus balucha balucha Uvarov

[Type as for species, above.] DIAGNOSIS. See iiey.

MEASUREJIENTS 31ale Females - so. So. Range 3Iean

Head width . . I 3 . 7 . 9 4 '6 -5 .6 5-13

Tegminallength . I 6.S . g 12.7-1g.S 16.87 Femur length . . I 9.1 . 9 I2*7-I7'4 Ij.06

Total length . . I 14.2 . 9 20.6-29.1 2j.00

i

- . .*u_-

A RE\-ISION O F T H E G E K U S C A L L Z P T A M U S S E R V I L L E 343

DISTRIBUTIOS. See Text-fig. 8, p. 304. Probably found throughout hüis of West Pakistan ; not below 1,000 m. A montane subspecies with intermediatec with subsp. brachykteriis (Dirsh) at Kabul River.

Baiuchistan, Up t0 3,300 m., 1 $, 8 9, vi-viii.

Easily separable from all sympatric species, e.5. C. ifnlicus (montane form), C. barbnnu bnvbartcs (orange-legged . form), C. coelesyvieiuis coelesyriensis (semi-desert form), and C. telzziicercis (extreme semi-desert form) (see key, p. 314).

hhTER+iL ESAMISED.

DISCUSSIOS.

)v. P-AKIST-4S :

Cnlliptamus balucha brachypterus (Dirsh) stat. n. Metvomerirs brachypterus Dirsh, 1937, Atti . Mics. Stov. nat. Tvieste, 21 (2) : 44. [Holotype 8, W.

Chitral reg., Birir, 2,300 m., viii.1955, ( A . Martrssi). Trieste Mus.] Pakistan,

DIAGKOSIS. See key, p. 342.

MEASUREMEXTS Males Females 1 P No. Range Mean No. Range Mean

Headwidth . . 8 3'6-4'0 3.81 25 4-6-6-1 5.26

Tegminal length . 8 j - 6 - g - I 7.60 z j 10.0-16.0 13-47 Femur length . . 8 9.2-10.8 9-84 2 j 13.2-16.9 15.25

Total length : . . 5 11.2-15-7 13.45 Z j 18.7-25.8 2 2 - 0 2

DISTRIBUTIOS. See Text-fig. 8, p. 304. Chitral and Kabul River systems up to

MATERIAL ESA;\IISED. AFGHANISTAN : Kabul, 1,600-2,400 m., 6 3, 22 9, .y.

2,400 m.

W. PAKISTAN : Chitral, 2,300 m., 3 8, viii.

Calliptamus coelesyriensis (Giglio-Tos), comb. n.

DIAGNOSIS 8. I . Lateral espansions of penis valves broad and auricular, hiding or partly hiding penis valves from view laterally (Test-fig. 16 : p. 317). Zygoma mith unique inflated posterior expansions (Test-fig. 16 : A, g). Dorsal ectophaiiic membrane plate with unique, erect sclerotization on median posterior edge (Text-fig. 16 : C, c).

2. Cercus with lower apical lobe usually undivided (Text-fig. 17: Fi, O. p. 319). Subsp. hissavicus however often has separate median apical lobe.

3. Tegmina with z or 3 branches of Rs. Koticeable concavity in membrane between CUI and CUZ (between CUlb and C U , ~ ) .

4. Melanic forms common. Buff tegmina peppered with darker pigment (like C. italicus in E. Iran), darker highland forms from Turkey and.Afghanistan with coarser spotting.

5. Inner side of posterior femora a pale shade of general body colour ; three separate brown spots (often faint, replaced by deep pink or absent), not extending on to upper side of femur. Lower inner carina and adjacent area pale ruby pink or greyish pink (Test-fig. 17 : P, Q, S, U. p. 319, cf. C. italiczts, Test-fig. 2).

6. Posterior tibiae some shade of pale rubv; intense on outer side. Infusions of melanic pigment gives mauve appearance.

7. Hind wings with pale ruby flush in whole of anal fan. 9. For differentiation froni C. ifnliciis see p. 314. If median femoral spot well developed, then

3.14 S . D . J A C O

posterior spot subequal, Text-fig. 17 : S. p. 319 ; in C. balrdia median spot always larger. Other colour characters as for male.

Similar t o C. teniricercis but with greater estension north eastwards (as subsp. hissnricz~s), see Text-figs. 8 and 9, pp. 304, 305. Does not enter Negev desert in Israel or N. Turkey.

DICCLXCIOS. For synonymy of JIetromenrs see introduction, p. 300. Subsp. nngzcstrts, defined partly on the narrowness of fastigium verticis in the male, cannot be separated on this character (see Text-fig. 17 : A-C. p. 319 showing range of variability from one locality). I t seems to be an upland form of the species with many facies of a montane form (see p. 304).

DICTRIBGTIOS.

KEY TO SUBSPECIES

,MALES

I . Penis valves flattened and appressed to cingular valve (Text-fig. 16 : D, B, m. p. 3 I 7) . Epiphallus (Test-fig. 14: G. p. 313) thickened uniformly in centre, without cruciform or narrowly parallel thickening (cf. Test-fig. 14 : A, B, D. p. 313)

coefesyriensis hissoricus Qíishchenko) (p. 345) KE. IRAN ; NE. AFGHANISTAS ; TADZHIKSKAYA REP., C.S.S.R.

-. Penis valves small, pointed (Text-fig. 16 : E, m), orientated with their flat surfaces in a plane parallel with long axis of body ; valves wholly or partly hidden from side by auricular accessory penis valve processes. Never with a median apical cercus lobe z

coelesyriensis coefesyriensis (ab. corbonaria Uvarov) (p. 344) 2. Black except for deep ruby-black tibia1 colour

-. Colouring light brown or buff with fine tegminal speckling, or darker brown with bolder tegminal markings. Epiphallus Text-fig. 14 : A, B, D. p. 309

coelesyriensis coefesyriensis (Giglio-Tos) (p. 345) ISRAEL ; JORDAh' VALLES ; SYRIA ; IRAQ ; I R A S ; s. TURKMESSKAYA REP., U.S.S.R.,

\V. AFGHASISTAS ; W. PAKISTAN ; TCRKEY.

Calliptarnus coelesyriensis coefesyriensis (Giglio-Tos), comb. n. Caloptenrrs coelesyriensis Giglio-Tos. 1893, Bid . Mzcs. Anat. comp. Tovino, VIII, 164 : 10-11.

[Holotype d, Syria. Turin Mus.] Calliptamrcs italicirs ab. carbonavin Uvarov; 1914, Rev. riisse Ent., 14 : 226. [Holotype 8,

Turkey, Ordubad prov., nr. Eriwan, I .vi.1911, ( K . A . Sntirvnin).] Kripa coelesyrieitsis angzrsfzrs Gvarov, 1934, Eos, 10 : IIS, Text-fig. ;Holotype 8, Turkey,

Ankara prov., Ankara, 1o.viii. 1931, (B. P. Uvnvov) Brit. bIus. (nat. Hist.l.1 Syn. n. CalliPtamzis fentiicevcis nnatolicics AIafan, 19j1, Sbov. ent. Odd. náv. ilízrs. Pnize, 27 : 384. [Holo-

type s. Turkey, Ankara Baraj, (3-4). vii. 1947. Sat . Aíus. Prague.] Syn. n. Metrotnems coslrsyriensis intvicntzis Nishchenko, 19 j1, in G. Bcé-Bienko S L. L. Mshchenko :

Opyed. Faum S.S.S.R., 40 : 261. [Holotype 5. Turkmeniya S.S.R., Oschek, Kopyet Dag, nr. Gollbeck, (21-23). v. 1913. Leningrad J1us.j Syn. n.

DIAGSOSIS s. Sre key. above. Sarrow zone of intergrading pliallic types with subsp. hissavicus

9. Use colour characters (see keyc, pp. 314 and above). (see Text-fig. S and Test-fig. IG : H, J).

A R E V I S I O N O F T H E G E N U S C A L L Z P T A i V U S S E R V I L L E 343

MEASUREMENTS Males Females -7 7-- ;?;o. Range Nean Xo. Range Mean

Femur length . . 82 9.0-14-3 11.54 209 12.1-21.0 16.12 Tegminallength . 84 S.&zo.4 13.56 212 11.6-27.4 18 .39

Head width : . - 87 3.5-4'7 4'53 214 4.2-6.0 5 .09

Total length . . 83 14.4-28-1 19.86 2 1 6 17.5-3S.I 27-14

DISTRIBUTION. See Text-fig. 8, p. 304, and discussion below. hL4TERIAL ESAJIISED. TURKEY : Ankara, 3 $, 3 $2, vii-viii; fhtalya, Up t0

1,900 m., 2 $, 36 9, vii ; Isparta, up to 1,700 m., g $, 19 9, vii ; Seyhan, up to 1,900 m., I $,4 9, ix ; Maras, up to 2,000 m., 2 $, I 9, vii ; blugla, up to 1,800 m., g 9, g 9, vii; Denizli, up to 1,900 m., 10 9, viii; Urfa, 4$, 6 9, vii. SYRIA: Damascus, I $, 6 9, vii ; Tekieh, 2 d, 3 9, vii ; Jebe1 Maza, I $2, vii ; En Kebk, I S, iv ; Aieppo, I 9, iv-vii. ISRAEL : Jerusaiem, I 8, v ; Southem distr., Beersheba, I 9, v. JORDAN : El Ghor, z 8, I 9, iv ; E. of Hebron. I 9, v ; Jerusalem to Jericho, I 3, 8 9, iv-v ; Khor Fasyii, 6 8, 4 $2, iv ; \Vadi Shueib, I $, 3 9, iv ; Wadi Keld, I $, I 9, v. IRAN : Dowom, 22 8, 19 $2, v-vii; Haftom, 7 6, 7 $2, v ; Markazi, 5 $, 13 $2, Mi-ix; Hashtom, 7 8, 8 9, iv-vi; Dahom, 2 8, I 9, vi. U.S.S.R. : Kazakhstan S.C.R., 3 9, ix ; Armeniya S.S.R., 7 $, 7 9, vi ; Turkmeniya S.S.R., I $, I 9, l-.

DISCUSSIOK. Two main forms occur, a pale buff semi-desert type with melanic forms (increasing in proportion north eastwards), and smalier, dark coloured upland forms which often display var. margiitellzis. The two forms intergrade over a broad zone.

Calliptamus coelesyriensis hissaricus (hIishchenko) comb. n. Metromerus coeleswiensis hissavicirs Mishchenko, 195 1, in Be6-Bienko and L. L. Mishchenko,

Opred. Fazrne S.S.S.R., 40 : 261. [Holotype 8, U.S.S.R., R. Khozov-RIer, Ober Eeskander- Komya, viii. 1947 (A. Kerichenko). Leningrad Mus.].

DIAGXOSIS. See key to subspecies, p. 344. Inner side of posterior femora usually deep blackish ruby to maroon. Inner femoral spots separate, median spot largest, often diffuse (Test-fig. 17 : T. P. 319).

MEASUREMESTS Males Fe males -7 - So. Range Mean No. Range Mean

5-00 Head width : . Femur length . . 11 7.7-11'8 10.73 6 10.3-1;.8 16-02 Tegminal length . 11 8.9-14.4 12-85 6 13.0-2j.g 21.13

6 4.1-j.~ . 11 2.5-4.1 3'75

Total length . . 11 14.7-20.2 18.66 G 1S..+--<4'4 29 .2s

DISTRIBUTIOS. See Text-fig. S, p. 304. Found in tributaries of h u Darya R. hystem, KE. Iran, and probably Golodnaya Step' eastwards (S. Turkmeniya, and Tadzhikstan, S.S.R.).

Badakhshan, Shiva and Senna, ~,Soo-z,Soo m., 4 8, I 9, vii. U.S.S.R. : Type and allo tjpe.

3LATERI.4L ESAJIISED. IRAS : Dovvom, Shahrud, 6 3, 4 0, v-vi. AFGH-4NISTAN :

346 S . D . J X C O

DISCUSSIOX. For resemblance to Imlomerzu see p. 297. The subspecies show a narrow belt of intermediates with the nominate subspecies in XE. Iran (see Text-fig. 16 : H, J. p. 317 for intermediate phaiiic types).

0.5 rn m.

1 ' r n /

FIG. 26. Apical portion of phallic comples in C. plebeius (Walkerj (.l. dorsal, B. lateral, and E. posterior aspects) and C. rizndeirne üvarov (C. lateral and D. dorsal aspects).

X R E V I S I O S O F T H E G E S G S C A L L I P T A i l f U S S E R V I L L E

Calliptamus plebeius (Walker, 1870) 347

- - Hcteracris plebeia \Valket;lSyo; CataGgiie of the Specirizeits of Dermaptera Saltatoria in the British

Mttsezwz, 4 : 673. [HoloQTe q, Canay I., Brit. Aíus. (nat. Hist.). Descriptions based on male plesiotype, Gran Canaria I., Fataga, viii. 1931 (H. B. Cott). Same depository.]

Caloplenits udcaiiizts Krauss, 1892, Zool. r4?lz.. 15 : 167. [Unknoivn s o . of syntypes &2, canary Is., Tenerife, Palma, Gran Canary, Hierro. Vienna BIus.]

DIAGSOSIC J. I. Genitalia rather membranous (Test-fig. 26 : A, B, and E. p. 346). Penis valves membranous, hidden by membranous cingular valve from above. Valves not hidden from side by lateral accessory processes ; cingular valve weakly sclerotised, with flat obliquely sloping posterior surface. Cingular arch covering base of aedeagus not constricted behind valves (cf. C. tenzticercis), its posterior edge rising vertically (not obliquely as in C . mudeirac,

2 . Cercus unique ; upper lobe very broad, lower apical lobes weak and divergent (Test-fig.

3. Tegmina always surpassing knees of folded posterior femora. 4. Colour polymorphism as C. barbarus but al1 forms much darker. 5. Inner side of posterior femora yellow to orange-red or red. Inner femoral spots always

fused. Hind femora narrow for genus (escept C . madeirue, where just as narrow), i.e. length t o depth ratio never less than 3-6 ; rest of genus 2-85-3-67.

P. 348).

24 : XIX. p. 336).

6. Posterior tibiae yellow to orange-red, to red. Spines rather elongate. 7. Hindwings colourless, hyaline, slightly infumate. 8. Lateral pronotal carinae straight, diverging uniformly backwards. Kever converging in

last 4 of their length. 0. As for male.

NEASUREMESTS híales Females P 7- Ko. Range Mean h'o. Range hlean

Head width . . 8 3.0-3-7 3'53 46 4.1-4.9 4.48 Femur length . . S 9-5-12.8 11.49 46 15*2-18.5 16.81 Tegminal length . S 13.1-18.0 14-90 46 21+6-25.7 23-94 Total length . . S 17.8-24.1 22.63 46 29.2-34-8 3i-30

?

DICTRIBCTIOX. Found on Gran Canaria, Hierro, and Tenerife in the Canary group, and probably other islands. Localities from sea leve1 to 4,000 ft.

Iv~ATERIAL ExAMIKED. GRAx CANARI.~ 1. : Fataga, Las Palmas, Caldera Bandana, 13 8, 56 9, Mii-ix.

HIERRO 1. : El Golfo, I $, viii. TESERIFE 1. : I 9, -. D~SCÜCCIONT Can live on very arid lava slopes among sparse vegetation. Shows

aii colour polymorphs over a very smaii area of distribution. Rfisidentified as Calop- felzus ifnliczrs by Heyden (1'872) and C. barbnrzis by Ramme (see p. 334 for reference to Willemse, 1936).

--- .--.

Calliptamus madeirae Uvarov, 1937 Calliptamzis madeirne Uvarov, 1937, Ark. Zool., 29, A(15) : 4. [Holotype 8, Madeira, Funchal, (27-30) . i s . 1910. Stockholm iiíus.]

DIAGSOSIS d. I. Lateral accessory processes of penis valves espanded, almost hiding penis valves from view laterally (Test-fig. 26 : C, D. p. 346). Cingular valve as in C. barbarus, not

t

348 N . D. J A G O

elongate, inflated, or membranous. Cingular arch unique, with two gently sloping lobes a t its posterior end (Test-fig. 26 : C, b and g), posterior edge of arch not being inflated in any way. 2. Cercus as in C. barbarus ; apical lobes separate. 3. Tegminal size as in C. plebeiris. 4. Colour forms as in C. plebeius and C. barbarzts. j. Inner femoral spots as in C. plebeitrs. Lower inner carina dull ruby-red, or buff body colour. 6. Posterior tibiae dull ruby-red, never yellow. 7. Hind wings suñused with pink on anal fan. 8. Pronotal morphology as in C . Plebeircs. 9. Diagnostic features as for males.

.x

MEASUREMENTS hfale Females P

Head width . 1 3’4 . 10 4’5-j.1 4‘73

SO. Ko. Range hl ean

Femurlength . I 10.3 . 10 16.1-19.6 17.57 Tegminallength . I 14’2 . 10 21-4-28.0 24-20 Totallength . I 19.6 . 10 29-5-37.4 32.63

DISTRIBUTION. MATERIAL EXAUIKED. MADEIRA 1. : Various localities up to 1,200 m., I 8, 7 9,

viii-x. DISCUSSION. The speciec may have evolved from C. barbarus due to chance

introduction from the Iberian peninsula, i.e. C. plebeius may have been denved similarly from N. Afnca.

Known only from hladeira Island.

REFERENCES

ADAMOVIC, Z. R., 1956, Grasshoppers Calliptamus italicus (L.) and Calliptamrrs barbarus (Costa) in south Banat, Serbia [In Serbian with English summary]. Zborn. Mutitse Srpskc, Novi Sud Ser. prirod. Nauka, 11 : 123-135, 4 figs.

ALEXANDROV, N. V., 1947, Les Acridiens des régions Nord, Sord-Est, et Nord Ouest de 1’Iran [In Russian with French summary]. Ent. Phytop. appl., Téhran, 3 : 61 j, I map.

BATRA, H. X., 1956, Certain pests of fruits likely to be introduced into India from North-West Frontier Province (West Pakistan). Indian J. Ent., 18 : 63-75.

BEE-BIENKO, G. YA., á- MISHCHEXKO, L. L., ~gjr, Acridoidea of the Fauna of the U.S.S.R. and adjoining countries [In Russian]. Opred. Fazcne S.S.S.R., Moscow. 38 : 1-378.

BOLIVAR, I., 1876, Sinopsis de les Ortóptéros de España y Portugal. Ann. Soc. esp. Hist. nat. 5 : 292, 296. - 1898, Catálogo sinoptico de los Ortopteros de la fauna Iberica. Ann. Sci. nat. Porto, 4 . BRUNNER vox WATTENWYL, C., 1882, Prodromzts der Ezwopaischen Orthopteren.

BURMEISTER, H., IS~S, Handbzcch der Entomoiogie 2(2) : 591-664, 1013-14. CHOPARD, L., 1922, Orthopthes et Dermaptkres, Fatcne de Frunce. Lechevalier, Paris. - 1943, Orthopteroides de 1’Afrique du Kord, Faune de I’Empire Francnis (1) : 1-450, 658,

CHETYRKISA, 1. A., 19 jS, The Italian Locust (Calliptamris italiczis (L.)) in Eastem Kazakhstan

CHORDBAZHIER, P., 1926~2, Injurious Grasshoppers in Bulgaria [In Bulgarian with French sum-

DIRSH, V. X, 1956, The phallic comples in Acridoidea (Orthoptera) in relation to tasonomy. Trans. R . ent. SOC. Loird., 108 (7) : -23-356, 66 PIS.

466 pp., 1-11. Leipzig.

figs .

[In Russian] Trrrd. usesoyziz. ent. Obshch., il-loscow, 46 : 5-67. I map, 3 figs.

maryj : Rev. Iizst. Rech. agron. B d g . 4, (1-2) : 169-181.

.A R E V I S I O S O F T H E G E S U S CdLLIPTAJIGS C E R Y I L L E 349

ESE, 11.. 19 36, Zwei für Uhdschutzstreifen schadliche Heuschreckenarten IIn Rumanian with German and Russian summaq-j : Rec. Padurilor, 71 : 113-11j, 6 figs.

FABER, X., 1949, Eine bisher unbekannte Xrt der Lauterzeugung europaischer Orthopteren : 3iandibellant \-on Cnlliptnnziis itnliczrs (L.) : Z. Snt t i r f . , 4b : 367-369.

FILIPJEV, i. S., 1626b, Plant Food. Injurious Insects and other Xnimals in U.S.S.R. in the years 1921-1924. So. 2 -1cridoidea [In Russian with Engiish summary] : Bur. appl. Eizf. St. It is t . exp. =Igroiz., Leiiingrad, 13 : 57-176. 10 maps, I fig.

- 1929, The Locust question in Soviet Rusia . Tvairs. 4th inf. Coitgv. Eut., Itka.cn, S.Y., 2 : So3-S12, I map.

FISOT, A., rSS3, Les Orthoptires de ln Frunce : 109 pp., I pl., Paris. - 1890, F m m e de In Frnnce, Insectes Orthopteres proprement dits : 322 pp., 13 pls., IS figs.,

FISCHER, L. H., 1Yj3, Ovfhoptern Etrropaea : 4 j 4 pp., IS pls., Leipzig. FISCHER DE \VALDHEIJI , G., 1S20-1S40, E~itomographie de la Rirssitz. Orthopteres de la Russie.

GRACSE, P. P. Ci. HOLLASDE, -1., 1945, Biological and Systematic notes of French species of the genus Calliptnmics Serville : .Irch. 2001. exp. gen. 84 : 49-69.

HOLZEL, E., 19j j, Heuschrecken und Grillen Karntens : Carinthin 11, Klngenfzirt Soizderheft 19 : I I Z pp., 2 col. pls., 2.1 figs.

JACOBSEX, G. G. CP: BIAXCHI, V. L., 1902, Orthofitera and Odonatn of the Riissian Empire and adjoining cotintries [In Russian]. : s?c -b 952 pp.

JANNONE, G., 1936. Kuovi contributi alla conoscenza della Fauna delle isole Italiane dell’Egeo T‘. Studio bio-ecologico e systematico dell’ Ortotterofauna con notizie sui Blattoidei, Jlantoidei, e Fasmoidei. Boll. Lab. Zool. Portici, 29 ; 47-245. - 1935. Primo contributo alla conoscenza dell’ Ortotterofauna della Libia. Boll. Lab. 2001.

Portici, 30 : 87-120, 5 figs. KIRBY, \V. F., ~ S g o , The employment of names proposed for genera of Orthoptera prior to

1S40. Sci. Proc. R. Dtiblin Soc. (N.S.) , 6 : 556-597. - 1910, A synonymic cntalogzre of the Orthoptera, Vol. 3, Orthoptern, Snltatovia. Part 11.

Loczrstidae cel Acridiidae. KIRICHESKO, A., 1926, A study of the ecolo,gy and biology of Cnlliptamus italiczcs (L.), in the

steppe zone of the Ukraine. [In Russian] Odessn Reg. Agric. E x p . Stat., Ent . Dept., 1 : 47

KOBAKHIDZE, D. S., 195 j , Injzrriozrs Entomofauna of Agriczrltziral Crops in the Georgian Soviet

LA GRECA, 31., 19 j&, Significata biogeografico di ripartizioni disgiunte in Ortotteri non montain

- 19 j9 , L’Ortotterofauna pugliese ed il su0 significato biogeografico : i l J e m Biogeogr. adriat.

LISSAECS, C., i j j S , Systemz Snti irne, 10th edn., 1 : 424-433. M A G A S , J., rgjI , Results of the zoological scientific espedition of the Sationai 3luseum in Prague

to Turkey. Sbor. eitt. Odd. nár. Mars. Praze, 27 : 59-64. - 1952, Calliptamzrs bnrbarzis Costa, a new kind of grasshopper for the fauna of Czechoslovakia

;In Czech!. Sbor. eitt. Odd. ndr Mus. Praze, 28 (406) : 149-1 j6 . BIATVEJER, A. & S., 10j6, On the spread of some species of submontane locusts according to

altitude zones [Iii Croatian with English summary]. Zasht. BilB, Belgrade. 33 : jj-SS. ~ ~ I S H C H E S K O , L. L., 19 37, Fnimn of tlie U.S.S.R. Acrididne (Catnizfopinne), OrthoPfern Hnl.

;In Russian:. 4 ( 2 ) : 610 pp. Leningrad. PESER, 31. P., 1960, The biology of Calliptniniis palnesfiizeizsis Bdhmr. with special reference to

the development of its eggs. Btrll. Res. Cotinc. of Isrnel, 9B (2-3) : 131-rj6. RADCLYFFE-ROBERTS, H., 19-11, -A comparative study of the subfamilies of the Xcrididae

(Orthoptera) primarily o11 tlie basis of their phallic structures. Proc. Acnd. imt. Sci. Philad. 93 : 201-246.

RACGE, D. R., ig5j , The wiiig-ceirntioi~ of the Orthoptern Snllatoria. 1j9 pp. 106 figs. London.

Paris.

37 PIS.

St. Petersburg

pp., 2 pls.

Socinlist Repiiblic [In Russian]. 263 pp. Orthoptera pp. j9-6s. Tiflis.

d’Italia : Arch. bot. biogeogr. itnl., Forli, 32 : 113-129, 3 maps.

4 : 58-99, figs. 103-104.

.J

t i

350 S . D. J ; \GO

RAhth iE , W., 19 jI, Zur Systematik Faunistik und Biologie der Orthopteren von Südost-Europa und 1-orderasien. .lIitt. rool. -1Izts. Berl. 27 : 1-431, 39 pls., 134 figs.

SERVILLE, J. G. A, 1S31 , Rewe Aiethodique des Insectes de I'Ordres des Orthopt6res. A n n . Sci. izct l . (Zool.) 22 : ?S-új, 134-162, and 261-292.

- 1Y3S (Dec.), Hidoive .Ynliirelle des Insectes In Roret, Collectiou des Suit?: k Bii.fion. OrtRopt&es: 776, p. 14, pls.

SILVESTRI, Ir., 1934, Compendio di Entomologia Xpplicata : Parte Specinle. Portici Stnb. Tip. Bellavista, 1 : 99-111, figs. S4-93.

S T ~ J . , C., 1873, Recensio Orthopterorzim, 1 : 1-154. STEBAEV, 1. V., 1957, The Orthopteran fauna in the landscape of the principal watershed in

N. Ergeni [In Russian with English summary]. Zool. Zh., Moscow, 36 : 396-407. 2 figs. TARBIXSKY, S., 1930, Contribution to the knowledge of the genus Cnlliptamirs Serville (Orthoptera)

[In Russian]. Bzill. Acad. Sci. U.R.S.S. (Cl. Sci. Plzw.-Alnth.), (2) : 177-186, 10 figs. CVAROV, B. P., 1922, Sotes on the Orthoptera in the British Museurn. 2 . The Group Calliptamini.

Tvans. ent. SOC. Lond. 1922 : 117-177, pl. I . - 1943, A revision of the genera Sphodromerus, dletromer~is, and Splrodroitotats. PYOC. Linn.

- 1950, The genus Caloptenopsis 1. Bolivar and its allies. Eos. Tomo extraovd., Bolivav

T-ARDÉ, V. P., 1934, The protrusible vesicles in Cyrtacanthacrinae, Acridiinae (Orthoptera).

VASIL'EV, Ii. A., 1g5oa, Migratory flights of the Italian Locust Calliptatnzrs italz'ctts (L.) [In

- 1g5ob, Phases in the Italian Locust (Calliptnmiis italiczis (L.)) [In Russian!. C. R. Acad.

VOCSELER, J., 1902, Beitrage zur faunistik und biologie der Orthopteren Algeriens und Tunisiens.

WALKER, F., 1870, Catalogue of the specimens of Dermaptera Saltntoria in tlte collection of tlte

SOC. Loid. , 1941-1942 : 65-85, pl., 3 figs.

Memorial : 385-413.

J. Univ. Bombay, 2 (5) : 53-57, 5 figc.

Russian]. C. R. Acad. Sci. U.R.S.S.. (.!V.S.), 74: 385-388, 2 figs.

S C ~ . U.R.S.S., (N.S.), 74 : 639-642, 2 figs.

Zool. Jb . (Syst.) 16 : 337-404, pls. 17, 18, figs.

British IClzrseum, (3) : 485-.j94, and (4) : 6oj-801.

A R E V I S I O S O F T H E CESUS CALLIPTAJIUS S E R V I L L E 325

'' Okbnensis " \vas created to differentiate long-winged forms of the species. An examination of Test-figure IS, p. 321, which show-a tegminaljfemoral relationships in Sorth African material, indicates that the species tends to produce shorter winged forms in Cyrenaica and Libya. It is interesting to note that the epiphalli of series from the Alpes JIaritimes and Libya resemble each other quite closely, intermediate forms in the Eastern Pyrenees having smalier lateral fenestrae and a larger median fenestra, the lower median fenestra often being occluded (cf. Test-figs. 14 : H and E. p. 313). In epiphaliic characters therefore, material from Libya and south-eastern France, forms two groups of populations relatively isolated from their neighbours to the west, but apparently similar to each other. The ratio of male femur length to head width shows a rise eastwards in K. Africa from Jlorocco to Libya, i.e. Ait Bou Guem- mez 2-97, hlascara 3.0S, Boghari 3.23, Slonta 3 2 3 . The Atlas mountain ratio is similar to that for populations in the Alpes Alaritimes foothills, i.e. Méounes 2-97, St. Maximin 2.91. Females show similar trends, populations in the Atlas mountains showing a ratio value of 3.18-3'39, montane forms of C. tarunicus, e.g. from Afghani- stan, Senna (1800 m.) having similar ratios, i.e. 3.29.

1. P

Calliptamus turanicus Tarbinsky, 1930 Calliptamzcs italicus var. watfenwylianus Jacobsen & Bianchi, 1902, Orthoptera and Odonata of

Calliptantrrs turaniczis Tarbinsky, 1930, Bull. Acad. Sci. U.R.S.S.: 184. [Holotype $, Tadzhik-

I . Penis valves, cingular valve, and lateral accessory processes of penis valves like thoseof C. waffenuyZinnzis (see diagnosis of that species, p. 311 and Test-fig. 13 : A. p. 320). Cingular rami diverge dorsally. Ectophallic membrane plate (Text-fig. 13 : A, c, and F. p. 311) with laminar median posterior tip, only slightly thickened and rugosely sculptured. Ectophallic plate much longer than wide.

2 . Cercus with upper and louer apical lobes almost equal in size and length (like Test-ng. 24 : J. p. 336). median apical lobe being more or less obliterated (especially in montane forms).

3. Tegmina moderately to well developed, Rs possessing 3 or 4 branches (3 brancnes in montane forms from south of species range).

4. General colouring pale brown with darker brown markings. Ko evidence .of estreme " marginellus " variety, pronotum usually a pale form of type D (Text-fig. 19. p. 322) mith more uniform colouring, or as type H. Tegniina with discrete brown spots, never reduced to mere peppering as in C. coelesjriensis nor developed into bold transverse bands.

5 . Dorsal spots of posterior femora never extend across upper inner carina on to inner face of femur (Text-fig. 20 : J. p. 324). Inner area pale buff except in montane forms where it may be almost completely filled with a suffuse brown pigment. Lower inner carina often pigmented with pale orange pigment, though often this is absent.

6. Posterior tibiae deeply to lightly pigmented with orange-red on their inner and ventral surfaces, often with a pale zone about 5 mm. from their prosimal end.

7. Hind wings boldly suffused with pink, this colour fading apically. Folded wings with apices surpassing knees of folded posterior fernora. Wings shorter in montane forms, so that ming apices leve1 with, or just short of, knees of folded femora, e.g. in XE. Afghanistan.

9. Bigger than fernales of C. italicus from the same area. Othenvise al1 other characters identical.

the Russian Empire : 317.

stan S.S.R., Golodnaya Step'. Leningrad Museum.]

DIAGXOSIS $.

C

326 S . D . J . \GO

ME.~SUREYESTS Males Females h - f \

S O . Range >lean S O . Range Nean Head width . . 8 4 '2-j -0 4 - 5 8 . 15 6'3-7'1 6.56

LVing length . . 8 14.2-24.4 20.66 . 15 '4.3-37.2 32.73 Femurlength . . s 12.4-1j.S 14.68 . 15 '0.7-2j.7 23.36

Total length . . 8 21-0-32.3 27.89 . I j 39.1-j0.I 44'19

DICTRIBCTIOS. In U.S.S.R. (Turkmenskaya, Uzbekskaya, Tadzhikskaya, Iíirghiz- kaya, Kazakhskaya) up to eastern end of northern slopes of Tien Shan ranges, and into the north facing valleys of hfghanistan up to about 2,000 metres above sea leve1 (see Text-fig. 7, p. 302).

JfATERIAL EsAJIISED. AFGHASICT-~S : Badakhshan, Senna, 1800 m., 2 8, 19, vii. U.S.S.R.: Kazakhstan S.S.R., I 3, S 9, i x ; Tadzhikistan S.C.R., 3 d, 4 9, vii ; Uzbekistan S.S.R., I 3, I 9, viii.

Probably this species has evolved quite recently, after geographical ceparation from C. watte.tzuyZin.tzzrs, from which it is derived. Its southernmost populations resemble C. wnttenzeylinnus very closely in all but the phallic characters. The high passes, dividing the Amu Darya river systems from the Kabul River sj-ctem to the south, offer a barrier t o its espansion into southern Xfghanistan.

DISCUSSIOX.

Calliptamus abbreviatus Ikonnikov, 1913 Cnlliptainiis abbreciatus Ikonnikov, 1913, Iiuznetsk : 21. [Holotype 5, L-.S.S.R., Primorskiy

Kray terr., Ozero Khanka, Kamen' Rybolov, Sikhote Alin. Leningrad Aluseum.] CaZliptamrcs sibirictis Vnukovsky, 1926, Mitt. iiiiinch. ent. Ges., 16 : gr . [Unknown number 3

syntypes, U.S.S.R., SW. Siberia, Bezirk Kamenj. Tornsk State Lniversity.]

DIAGSOSIS d. I. Phallic complex like smaller version of C. itnliczis (Test-fig. 10 : A, B. p. 307). with penis valves relatively shorter. Penis valves project beyond tips of lateral penis valve appendices a distance equal to, or less than, length of lateral penis appendices themselves (Test-fig. 10 : B, m and 1). 2. Cercus apes variable. Usually like Test-@ 17 : H. p. 319. Lower pair of lobes may fuse

(but are always shorter than upper lobe, cf. siibdpinzrs, p. 338), or lower lobe may be very weakly developed forming a blunt obliquely rounded area below median lobe (as in some Stobbea

3. Tegmina tapered in apical z/3, when folded their apices falling short of knees of folded posterior femora.

4, Pronotal and body colour usually dark brown, speckled with blackjsh brown. Pronotum typically type E (Test-fig. 19, p. 322) ; little colour polymorphism. Tendency to form " mavgi- neilus " variety slight. j. Posterior femora pale body colour on inner surface, with three separate dark brown spots.

Lower inner carina and inter-carinal area sunused with dull ruby-red. Femoral spots variable in development, middle spot sometimes produced slightly below upper inner carina, while posterior spot reniains large enough to reach lower inner carina. Niddlc and posterior spots sub-equal in size.

SPP.1.

6. Posterior tibiae dull doep ruby-red. ;. Hind wings colourless. SeL-er so reduced as to be non-functional for tlight. $. 1-ery difficult to differentiate from C. s:ibnIpii~ris escept on distribution (see Test-fig. 6,

p. -;@r). Lnlike short-winged C. i f d i c r r s in .ifghanistan, never has pink hind wings. Other characters as males.

i

A R E V I S I O S OF T H E GEN'L'S C . 4 L L I P T i i d f C'S S E R V I L L E 327

~IEASUREJIESTS Jlales Females A A r -l r \

S O . Range Mean s o . Range JIean

Head width . . 19 3.0-4.1 3. jS . 16 3.3-6 O 4.235

Tegminal length . 19 7'7-11'7 9 ' j 7 . 16 r3.S-18.4 16.2j Total length . . 19 12.3-17.6 14*Sj . 16 19.6-30.4 24.23

Femur length . . 10 S.7-11.3 q.91 . 16 13-2-17'2 I j . j +

7

1

DISTRIBCTIOS. Present in northern China (Test-fig. 7, p. 302) , it estends into BIanchuria, northern Mongolia, north-western Kazakhctan, and into the south- eactern edge of the Siberian plateau.

J~ATERIAL ESAMISED. U.S.S.R. : Kazakhstan S.S.R., 6 8, z 9, vii-viii ; Karel' Skaya A.S.S.R., 2 9, s ; -4ltayskiy Kray, I 6, I 9, viii ; Siberia, 3 d, 2 9, viii ; Primorskiy Iíray, I s, I 9, is. C H I S A : Shantung, I s, I 9, is ; Heilungkiang, I I 9, - ; Hopeh, I 8, I 9, viii ; Gungulin, I 3, I 9, viii ; Iíiangsu, 2 3, z 9, vii-si ; Inner Jiongolia, 2 9, -.

DISCUSSIOS. Closely related to C. ifalicris (see p. 292 on probable origins), it is adapted to the \vide range of climatic conditions offered by the contrast between northern China and southern Siberia. Its reported distribution in the blediterranean region mas based on confusion with C. stibalpiniis. C. nbbreviatzis siciline has now been raised to species status, while it is clear that synonymy of C. icterictis with C. rzbbveviatiis by Tarbinsky (1930) was based on misleading euidence.

Calliptamus barbarus (Costa, 1836)

Synonymy under subspecies.

DIAGSOSIS s. I. Penis valves strongly sclerotized (Text-fig. ?j : A, D and F. p. 339), thickened and rugose or rather sniootherand elongate, wider dorsally (shown in T.S., Test-fig. 10 : E, m. p. 307). and often splayed out to give a spatulate outer margin to each (Test-fig. 2 j : J. p. 339). Lateral appendices of penis valves (Test-fig. 2j : D, 1) blunt, sclerotized, but never expanded in such a \va?- as to hide penis valves when viewed from a lateral aspect (Test-fig. 13 : D, f) . Cingular valve not usually heavily sclerotized, never overlapping penis valves so as to hide them from above (Test-fig. 2 j : D. m). Dorsal ectophallic plate with simple laminar, iipcurved, slightly thickened, posterior median lobe. Epiphallus very variable.

2. Cercus ve- variable (Test-fig. 24 : III-V and \-II-XVI. p. 336) . Lobes distinct, except in montane or boreal forms, where median and lower lobes merge (Text-fig. 24 : VIII. XI. XIV). Median lobe equal to, shorter than, or longer than lower apical lobe (if shorter then like C . sub- alpinns, Test-fig. 24 : XVIII).

3. Folded tegmina with apices surpassing (4 branches of Rs), leve1 with, or not reaching knees of folded posterior femora. JIay be short and functionless for flight.

4. Colour polymorphisni (see pp. 301-306) ver?- marked, especially in south and south-east of geographical distribution. " MnvgiiLeZlrts " form produced frequently. S o melanic forms. j. Posterior femora v e n variable, inner side L-aqing from pale dull yello\\- to orange or

bright crimson, with al1 possibie iritermediate shades. Iiiner femoral spots showing al1 degrees betweeii complete separatioii to complete fusion (Test-fig. 2 : B-D and E.'-H). Boreal forms often retain only median spots (formed by reduction of solid ovoid iiiner femoral spot found in southern populations). Fusion of spots initially betmeeii anterior pair of spots. Escessive deposi- tion of melanins leads to estension of inner femoral spot across lower inner carina, \\-here mised with red pigment a violet effect is produced (really a darlz ruby-red containing no blue element).

,

32s S . D. J A C O

Median inner femoral spot usually larger than other two, though in southern France often paralleling C. ifalicrrs very closely (Test-fig. 2 : E and H. p. 293). 6. Posterior tibiae pale dull yellow, to orange, to ruby-red. Black pigment produces dark

ruby colour, or dull pink. dependiiig on intensity of red component. 7. Hind wings usually strongly flushed with pink in at least basa1 half. Extreme montane

forms often have colourless wings. S. Tegmina usually coarsely spotted, but spot pattern ver?. variable. In “ nzarginellzis ”

variety wing may be uniform brown ; with pronotum t?lpe H (Text-fig. 19, p. 322) tegminal blotches almost disappear. Blotchec may va- in intensity, being sharply defined in semi-desert and yellow-legged forms (e.g. old f . pallidipes). Ksually parallels C. ifaliciis very closely where their ranges cross. Spot positionc not fised in relation to wing veins ; may be orientated at random or in roughly parallel bands across tegmen.

9. Very often difficult to separate from sympatric C. italiczrs, C. siibalpinirs, and C . uialten- vyZiaizirs (see p. 318, 9 diagnosis for italicus). Tegmina are not as abruptly tapered in their apical 2 /3 as those of szrbdpinrrs and wnfieitw]tlialzzrs in such localities. Yellow or orange-legged females possessing a single large inner femoral spot fiiling inner inter-carinal area can oniy belong to C. barbarus or C . tenuicercis. If orange-legged, then presence of orange colouring just posterior to black inter-carinai spot is diagnostic of C. barbarus. Female characters other than colour very uniform throughout genus. Maie material necessary for accurate determination.

~IEASCREJIENTS. See under subspecies.

DICTRIBUTIOS. \Vide geographicai range (Text-fig. 23, p. 333) ; occupying more southerly areas than C. italiccus (see Text-fig. 6, p. 301). Does not estend as far north as that species, e.g. in Massif Central of France it only survives in sheltered hoiiows and vdeys, and does not reach northern edge of the massif. In Korth Africa aititudes of 2,ooometresare attained, in Afghanistanmaterial beingcollected from 2,300metres. In southem Europe specimens seldom found above 500 metres (probably too wet). Occupies almost every dry Mediterranean or semi-desert region and steppe of the Palaearctic, including the islands of the Mediterranean.

~ L ~ T E R I A L ESAMISED. See under subspecies. DICCVCSIOS. Sow included in C. barbarus bnrbarzrs are subspecies and species

defined principally on colour characters, e.g. yellow or orange-legged forms include Cnloptesrtis discoidnlis, C. bnrbnrus var. pnllidipes, C. barbnriis i~tonspelliettsis, C. barbariis pallidipes f. snlilzn, C. cephalotes, Caloptemis itnlictis var. deserticola (last two identical) ; ruby-legged forms C. barbarus miniwms, C. barbnriis pnrvzis (both smaii boreal forms). Throughout the entire range of distribution the male genitalia offer uniform facies, except in Israel (see subsp. palaesti.nensis, Test-fig. 25 : J. p. 339). Elsewhere penis valve variation as a whole does not exceed that for any one locality. The folloning is a brief résumé of colour variation of this species throughout its range :

(i) ,Spain and Portugal. Ruby-legged in C. and S. Cpain, and Portugal (pale pink) : femoral spots bold and separate ; te,minal apices leve1 n-ith or surpassing knees of posterior femora. Pale orange-legged in S. and E. Cpain : single black inner femoral spot. IWng length as in firct form. Probably narrow zone of intermediates.

(ii) France. Ríassif Central and Ile d’Oleron series, crnall sized ; ruby-legged with separate inner femoral spots. Provence series similar ; “ ))inrginelltis ” form commoner near coast ; tegmina fa11 short of knees of posterior femora in series above 400 metres. Coastal forms have median femoral spot enlarged ; upland or

b

U

A R E V I S I O S O F T H E G E S U S C A L L i P T A J l U . 5 S E R V I L L E 329

boreal forms spots subequal. Delta and dune populations with fused femoral spots (former subcp. monspelliensis).

(iii) Italy. Xorth-south cline. In north like France. Coastal forms tend to have te,smina surpassing knees of folded posterior femora, but in C. Italy may vary in one locality from falling short to surpassing knees of posterior femora. In S. Italy long-winged form predominates, except at higher altitudes, e.g. ZOO-jOO m. near Cocenza, where al1 have tegminal apices falling short of knees of posterior femora. S. Italy all forms orange-legged (gradual cline from N. Italy) ; middle inner femoral spot enlarges (however reversed at high altitudes, e.g. Potenza 850 m., where red- legged with sub-equal spots).

(iv) Sardinia, Mallorca, Lipari 1s. Orange-legged. Tegminal apicec tend to surpass knees of posterior femora (except in Sardinia).

(v) Greece. Sorth-south cline from red to orange to yellow-legged forms (i.e. further east and drier). NE. Greece series orange-legged ; spots fused or separate ; like W. Turkey. From about Trikkala southwards, wings longer and legs paler orange (see sexual polymorphism, p. 303). Middle spot enlarges. Pelópormisos series all yellow-legged, folded tegminal apices surpassing knees of postenor femora (reversal of trends on Mt. Kelmos, grey-ruby legged at goo-r,ooo m.). Nauplion series show males ranging from orange-legged to yellow-legged. In S. Greece forms with single fused femoral spot predominate.

(vi) Crete, Sporádhes, Cyprus. In Cyprus yellow-legged (escept llt. Troodos, see p. 303) ; single fused inner femoral spot. In both other islands bright orange- legged series ; dark brown " inargiirellzis '' forms ; in Crete many specimens show separate inner femoral spots.

3 0' 3 5 O 40' 4 5 O 50' 55' 60' 65' 70'

Isoliyets (nicnii nnnrial rainfall in inchec) for 3Iiddle East (various sources). FIG. 2 I .

. .-

a

U

330 S . D. J . \CO

B

orange&ellow gre&dull r u f l inner fem&biae ruby oronge re femoral

spots

Fxc. E. C. bnrbarzis (Costa) (-4) and C. leizuicsrcis Tarb. (B) : dictribution of posterior femoral and tibia1 colour forms in Middle East and central Asia. Iíey to symbols for colour and melanic inner femoral spots below Fig. B.

I

?

33 * -\ R E I ’ I S I O S O F T H E GEXCTS C A L L I P T A 3 f C . S C E R V I L L E

(vii) Turkey. Four main colour types which intergrade : (n) SmaU ruby-legged form ; separate femoral spots, median largest : range

SIY. Turkey and al1 S. Turke- (identical with C. France, Kashmir, and SE. Greece material).

(b) Small yellow-legged ; separate to completely fused femoral spots. C. Turkey (Ankara-.4masya plateau) .

(c) Orange-red legged forms ; larger size than (a) or ( b ) . Very polymorphic (like Crete and Sporádhes material). -3bove 3000 ft. inner femoral spots separate (like (a)). iJ’. and SIY. Turkey.

(d) Large orange-legged form ; yeliow-legged forms present nith al1 grades to orange form. Inner femoral spots always a fused block. Wing apices exceed kneec of posterior femora. SE. Turkey into Syria.

The remaining populations of C. barbarzis can be split into 4 clines : (o) North Africa. Extreme north of Morocco has yellow-legged populations (inner

femoral spots separate to single solid marking.) Transition over narrow zone to orange-legged form ; tegminal apices leve1 with, or surpass, knees of folded femora. I n eastem Algena, Tunisia and Libya, single black inner femoral spot extends below lower inner carina, and darkens orange-red pigment below. I n upland areas, e.g. Chrea, nr. Blida (13,oo m.), replaced by smaller red-legged form vith separate inner femoral spots, this montane form reaching extreme of brachq-pterism in Atlas mountains.

(b) Bulgarian-Caspian cline. Gradual change eastwards from smali red-legged, short-winged form, to a longer-winged, orange-legged form, whose inner femoral spots are completely fused. Transition complete in region of Caspian Sea. Popula- tionc in Kazakhctan and Tien Shan very uniform, inseparable from C. b. deserticola.

(c) Kazakhstan-IYest Pakistan. Almost completely uniform ; orange-legged with fused inner femoral spots. South of high passes of Hindu Iíush, black area extends below lower inner carina [as in N. African material above), producing blackish ruby-red colour. Thus in contrast, material from Lake Balkash region has only orange on lower inner femoral area. I J h g length decreases with altitude up Amu D a n a river system, eventually tegminal apices in ail specimens falling short of knees of folded femora. If in a lowland population, tegminal length varies so that in some, tegminal apices surpass knees of posterior femora, while in others they fall short of knees of posterior femora : long winged type is always found among larger insects. East of Chitrai, Iíashmir populations are red-legged (cf. X. Turkey).

(d) Iranian- Jordan rift valley cline. Orange-legged, semi-desert form found throughout Iran (south of Elburz mountains), Iraq, and Cyria. Black area developed masimally in S. Iraq and SW. Iran. I n Jordan orange-legged form dominant on east side of Jordan valley westwards forrning transition to yellon--1egged members of nominate subspecies and finally yeilow-legged subsp. pnlnestinensis (Text-fig. g, p. 303). Orange-legged form in Segev desert, and at Iskerderim and Yerevan (Turkcy) has transitional and yellow-legged forms sympatric with it.

Yellow-legged C. temiicercis and C. bavbnviis are often difficult to separate (refer to Test-fig. 3 2 : -4 and B, p. 330). In Turkey both sexes of C. ieitiricercis have separate inner femoral spots, \\-hile C. bnvbnrus often chows fusion (best therefore separated

1

z

33' S . D. J . 4 G O

on male genitalia, see key, p. 306, and Text-figs. I j and 25). In Syria yelloív-legged C. barbarzis has solid inner femoral spot and te,gminal apices which exceed tips of folded posterior femora. Sympatric C. tesiziicercis has separate to partly fused femoral spots, and tegminal apices which are leve1 with knees of posterior femora. In Israel both species have yellow-legged forms with solid inner marking, t u t this form of C . tenuicercis only sjmpatric with orange-legged forms (for separation see p. 328, 9 diagnosis).

The type of C. srzontanzis has been found to have genitalia identical with C. barbarus barbarus, C. ictericzis is synonymized with C. barbarus, as the descnption of the female type (type lost) which came from Cadiz, fitc known orange and yellow legged C. barbarus material from S. Spain. U'nfortunately the specimen was already badly broken when described, and probably faded, the description leading to the report of this species from Korth Africa and elsewhere. C. xattemuylianiis has been the species most often given this name (e.g. Brunner, 1882 ; Finot, 1SS3). C. siibalpinus was also formerly called " ictericrcs ". Grassé SC Hollande (1945) introduced C. ictericzcs chopardi to embrace pink-winged forms of C. subalpinzts, unfortunately choosjng a ruby-legged boreal form of C. barbarus as type. Synonymy of C. ictericzis tvith C. abbreviatzis (see discussion p. 327 under C. abbreviatzrs), led Grassé & Hollande to give C. icterictrs chopnrdi a very \vide range in southern Europe, the geographical range of C. abbreviatzis being added as well. C . barbarus ntinintzis and C. barbarus parvzts are small boreal forms of the nominate subspecies, C. barbarus stanzis being a montane form of C. barbarus cephalotes (for discussion of montane forms, see p. 302). Calopteno+sis pztnctata has genitalia identical with nominate subspecies ; Caloptenzcs discoidalis is a female type (not male as stated in description) of C. barbarlis barbarus (yellow-legged form, but outside geographical range of subsp. pakzestinensis).

C. palaestineitsis survives as a subspecies of C. barbarlis with unique genitalia (Text- fig. zj : J. p. 339). C. palaestiitensis erythrocrzernis is a montane form of it which falls as a sj-nonym.

Loss of the type of C. barbariis has required desi,pation of a neotype ; this has been chosen from material corresponding to the original description and from near the type localitj- in Italy.

'

KEY TO SUBSPECIES

1. Penis valves elongate, rather thinly sclerotized, with widely divergent dorso-laterally splayed flanges (Test-fig. 25 : J, m. p. 339). Cercus in high montane forms with tendency to weak differentiation of median and lower lobes. Tips of folded tegmina may just surpass knees of folded posterior femora. Posterior tibia1 colour and inner femoral flush, pale yellow to bright red. Inner femoral spots fused or discrete

barbarus paíaestinensis Ramme (p. 33 j) (LEBASOS RAXGES : \vEST O F DEAD SEA-JORDAN VALLEY, generally iii area with

mean annual rainfall 28 in. (700 mm.) or more). -. Penis valves blunt, rather strongly sclerotized. parallel ; slightly spla-ed on dorso-

lateral edges (Test-fig. zj : B. p. 339). Cercus usually with separate lower apical lobes (fused in montane forms). Tips of folded tegmina not reaching, to surpassing, knees of folded posterior femora. Posterior femora dull pale yellow, orange or red internally. Black pigment belom lower inner carina may cause orange-red pigment to assume niauve appearance . . barbarus barbarus (Costa) (p. 333)

h

A R E V I S I O K O F T H E G E S U S CALLfPTAJfUS S E R V I L L E

Calliptamus barbarus barbarus (Costa)

333

Acridiztnz bnrbariim Costa, 1S36, Fairnn del Regno di .Yapoli. Ovfoffevi : 13, pl. I. -I-D, Saples. [Type lost. Seotype s, Italy, Salentina pen., Lecce prov., S. Puglie reg., Maglie, :o m., 2g.is. 1937 ( F . E . Zecinev). Brit. Jlus. (Sat. Hist.):.

Calopfeiiirs siczrliis Burmeister, 1838, Hnndbiich dev Eiitoriidogie. 2 ( 2 ) : 639 ; Ramme, I9 j I : Jl i t t . 2001. M a s . Berl., 27 : 311. [Type lost.:

Calliptamrrs ictericzrs Serville, I S38, Histoire .Yntirrelics des Insectes. In Roret, Collection des Szcites h Birffon. Orfhopt2res : 689, Paris. [Holotype 0, Spain nr. Cadiz. Type lost.] Syn. n.

Calliptamzu cephalotes Fischer de TYaldheini, 1846. Enfomogrnphie de la Rirssie, 4 : 243, hloscow. [Types lost.]

Caloptenus discoidalis IYalker. 18 70 , Cntalogirc of the Specimens of Dermapteva Saltatoria in the Collectioii of ihe Bvitish Mzrseirnz, 4 : úS4, 656 ; Uvarov, 1922 Trans. ent. Soc. Lotid.: 136. [Holotype (not s), Lower Egypt, Brit. Ríus. (nat. Hist.)] Syn. n.

Calliptamzrs miniwttrs Ivanov, 1888, -4rb. Ges. A-ntuv. f v . c'niv. Claarkov, 21 : 35. Syn. n. Caioptentrs itnliczis var. deserticola Vosseler, 1902, Zool. Jb . (Syst.), 16 : 395 ; Chopard, 1943 :

Fatrne Einp.fvanq., 1 : 404. [Syntypes JQ, Tunisia, nr. Gabes, 1902 ( A . Weiss).] Syn. n. Cnloptenopsis piinctata Kirby, 1914, Tlie Fntrnn of Bvitisli India including Ceylon and Burma.

Orthoptera (-4crididae) : 260, London. [Syntypes dQ, W. Pakistan. Jammu and ICashmir state, Baltistan reg., Brit. Mus. (nat. Hist.).] Syn. n.

Calliptamus montanus Chopard, 1936, Bull. Soc. Sci. nat. dlaroc, 16 : I 77. [Holotype d. Morocco, Moyen Atlas, Afraou des Beni Abdallah, 2,5 j o m. Paris Mus.] Syn. n.

Calliptamzrs bavbartu nionspelliensis Grassé & Hollande, 1943, Avch. Zool. exp. gén., 84 : 49-69. [Unknown number of syntypes, JQ. Paris RIus.] Syn. n.

Calliptamzcs ictericzis chopardi Grassé & Hollande, i g t j , Arch. 2001. exp. gén., 84 : 49-69. [Holo- type $., France, Fontainebleau. Paris Mus.] Syn. n.

Calliptamus barbarus paliidipes Ramme, i g j i , Mitt. tool . .iIirs. Berl., 27 : 311. plorocco. Paris Mus.] Syn. n.

Calliptamirs barbnriis nanzcs hlishchenko, 1951, In G. G. Be;-Bienko & L. L. Rlishchenko, Opred. Faune U.S.S.R., 38 : 257, Text-figs. [Holotype ,3, Tadzhikistan S.S.R., Zeravshanskiy Khrebet mts., Pendzhikent, Dolina Aranpaia. 1,500-2,050 m., 21. vii. 1942, (JI. Rzrbzcov). Leningrad Bíus.] Syn. n.

Calliptamtis barbarzrs pallidipes f. salina Maran, 19-54. Sbor. ent. Odd. nár. Mtrs. Praze, 28 (406) : 153. [Holotype J, Bulgaria, Sv. T'las, 6.viii. 1935, (L . Hoberlandt) Prague nat. 3lus.j

DIAGNOSIS. Cee key p. 332, and diagnosis of species p. 32 j .

FIG. 23. C. barbarzrs (Costa) ; trans-Palaearctic distnbution.

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