PRIRÜDOSLOVNI MUZEJ SLOVENIJE MUSEUM HISTORIAE NATURALIS SLOVENIAE SCOPOLIA Revija Prirodoslovnega muzeja Slovenije Journal of the Slovenian Mnieum of !\atural History^ 4911 CODEN SCPLEK • ISSN 035]-0077 öCuruijiA 4911 ülasilu PriTOdMlovnega muzeja Slovenije, Ljubljana Jfmrnai t}fihv Skivaniati Musntm of Natura! HisUny, Ljublfcinti Izdajatelj I Ediii'J by\ rrimdoslovni niuzej Slovenije, Ljubljana, Slovenija / StoveHian Mufieum of Natural Hijstdfy. Ijuhljuna, Slove tin Sofinanciratif' Subsidised by: Ministrstvo za kulluro in Ministrstvo za šolstvo, znanost in Spori Mimsiry of Culture und Ministri' of Education. Science anU Sporl Urednik / Editor: Janci; Gregori Uredniški odtor / Editorial Staff:_ Breda CtMC-]UHA>iT,[Huw GRIFFITHS (GSj Boris ICRYStUFRK. Lojtc MARIl^tEK. Ignac SIVEC, Kazimir TARMAN, Mikola TVRTKOVIČ (HR), Tone WRABER Naslov uredništva in uprave / Address of the Ediiorial Office and ^^dniinisiralion: Prirodoslovni mu/cj Slovenije, Prešernova 20, p.p. 290, SJ - 1001 Ljubljana, Slovenija Slovenitin A^Meum of Natural /fcton'. Prešernova 20. p.p. 290. J/ - HiOl Ljahljana, Slovenia TckoCi raCun pri LB / Current account at LB: 50100-603^40115 Lektor (za slovenSCino) i Reader {for Siowne): CvetanaTAVZES Lektor (za angleSifiino) / Reader (for English): Henrik CIGLIČ Oblikovanje ■' De.vijfn: Jurij KÜC BEK Tisk I Prinlatlhy: Schwarz d.o.o., Ljubljana l^ideta najmanj dve Številki lelno, niiklada po 600 izvOttüv The Journal appears a! least twice a }'ear, 600 copies per issue Natisnjeno / Printed: november 2002 Fotografija na naslovnici I Front cover: Jajce nove vrs(e Perlo iz Slovenije in Avstrije / E}fg of new spticies of Perlafmm Shvcnia and AiLttria. Revija je v podatkovnih ba^ah ' Journal is mvered by: COBIB. BIOSIS Zoological Record S™poliaNO49; 1-33(2002) The Species of Peria (Plecoptem: Perlidae): Evidence from Egg Morphology Ignac SIVEC and Bill P. STARK^ UDC (UDK) 595.735:591.4:591.9(045)=20 ABSTR.4CT The genus Peria is reviewed with a scanning dectroii micrograph study of egg morphology. Twelve species are recognized and placed in six species groups. Perln h(/rva!i, sp. tiov. and P. zwicki, sp. nov. are described from Turkey and abbreMata K.laj'Alek. and P. persica Zweck, are placed as synonyms of P. caucasica Glerjn. Key words: Plecoptera, Pcrhdae, Peria, taxonomy, egg morphology, description of new specics. IZVLEČEK Vrste rodu Peria (Ptecoptera: Pcriidae): doloiljivnst p« strukturi jajc.-Predstavljamo vrste rodu Perta na podlagi študije strukture jajc z elektronskim rastrski m mikroskopom. Ugotovili smo 12 vrst, kijih uvrščamo v Sest skupin vrst. Peria horvati, sp. nov. and P. zwicki, sp, nov. sta za znanost novi vrsti, ki ju opisujemo iz TurCije, P. abhreviata KlapAlrk and P perska Zwick sta sinonima vrste P. caucasica Gueajk, Ključne besede: Plecopterči, Perlidae, Perh, sistematika, struktura povrSine jajc, opis novih vrst. Peria, the oldest generic name still in use for a group of stonefly species, was proposed by Geokf^roy (1762) with the species type designated nt P. hipuneiata (Pictet) in 1963 through an opinion from the International Commission on Zoological Nomenclature. By the time of Illies {1966) catalogue this venerable name still retained considerable systematic baggage in the form of 293 species names, the majority of them listed as synonyms or as "Unsichere Arten", but also included were many poorly studied Asian species belonging to KamimuHa and other genera. SivF.r et al. (1988) accepted only eight species found generally from Britain, through the circum-Mediterranean region of Europe and North Africa to the Caucasus, hut subsequent studies (MtMBiELA 1990; Sivic & Graf pers.com.) support addition of two species not recognizcd in that study. 'Slcivcniai! Mitseum of Nstural History, t^jybljma, Slavenia ■Mississippi Collect;, Clinton, Mii^sis^sippi, USA Address torrespondence In: I SKet, Slovenian MiiSjCum üf Natural Hiilüiy, Preäertiova 20, lÜOt) Ljubljana. Slovenia. The geiius has never been studiL^ comprehensively and mosi speeies determtnations follow ttie works of AußbkT {1959), DtsPAX (1951) and Ili.ihs {] 955) each of which included the same four species. As other valid species were added, supporting data for their recognition was given (e.g. Bra.\sch & Joost 1971: Bhrthelemy & Terka 1980; Z wick 1976) creating a inosaic array of literature needed for species determination, and bccausc the majority of specimens coMcetcd are lar\'ae or females, the delermination of thase, based on a few characters variable over a wide geographical area, has become quite unreUable. We began acquiring egg material for a survey of ail known Per la species in the mid 1980's and continued the intermittent study of this material wilh scanning electron microscopy throughout the 1990's. Our objectives wene, 1) to discover egg characters which permit reliable detemiinarton of females and mature female larvae tliat contain eggs, and 2) to utilize these characters to gain a preliminary understanding of spec ics groupings within the genus. The results given below support recognition of a minimum of 12 spccics, including two previously uiidescribcd; these are p lac cd in six species groups, The results also indicate significant polymorphism exists in R pallida Gufrfn and P. marginufa (Panzer) which will require additional study. Methods for this study are given in Sivec ei aL (19S8), Specinnens used were made available frorn the following museums and individuals. Abbreviations are used with specimen data to indicate the sources of material. Bill P. Stark. Clinton, Mississippi (BPS), Biologische Station, Umz (BSL), Lminologische Flussstation, Schlitz (LPS), Monte L, Bean Museum, Brigham Young University, Provo(DYU), Musee Zoologique, Lausanne (MZ), National Museum of Ireland. Dublin (NMl). National Museum ofNatural Histor>', Prague (NMP). National Museum, Skopje (NMS), Slovenian Museum of Natural History, Ljubljana (SMNH), Stanley \V. Szczytko, Stevens Point, Wisconsin (SWS), Zoological Institute, Russian Acadcmy of Sciences, St, Petersburg (71SP). TIil- SptCKi 4lf ffWa ffletiipleni; Perlidai): l-vidci!« froir lijji Murpliclajj The PeHa hipunctata Group Perla hipunctata Pectet^ 1833 Figs, Egg,- Length 0.45-0.47 mm, width 0.2741.31 mm. Collar short, about O.tll mm long and about 0,09-0,12 mm wide. Circumference of collar with about IS irregularly spaced, obscure vertical ribs extending from rim onto shoulder» of egg body; a single pit located between ribs at collar base. Rim relatively smooth and without indsioris. Churionic follicle ccll impressions conspicuous and with center of each cell depressed creating a coarsely pitted appearance over the entire surtacc. Micropylar orifices moderately large with thin rim and vertical canal ridges extending about 0.03-0,04 mm from orifices; micropylar row sel about 0,13 mm from pole. SEM Specimen Data.- Belgium, Streupa. Jjne 1936 (MZ), Ireland, K.iHarney Kelly, Tore Stream. 30 May 1995, O'Connor (NMI). Scotland, Carluke (MMP), Distributioriv- Reported from Grsial Britain to the Balkans and North Africa. The only confirmed specimens we have seen are those listed above. Many specimens determiiied as P. bipurictata in European collectionsi, iin;luding several of those tram Morocco, are P. pallida^ but figures in Despax (1951) of male genitaliu and in Berthe^lümy Laur 0975) of eggs suggests populations of the species may remain in the Massif Central of France. Comments.- The type of bipunctata from Switzerland is lost (ZwiCK 1972) but the nanne has been applied rather consistently Co a species which may never have occured there. We are unable to locate specimens of P. hipmctata (in the sense of Ii.l ies 1955 and other authors) from mainland Europe, except for the possible specimen we list above from Belgium or the possible speciinens of BüHTHElemv & Lalr (1975) or D[y>pa\ (1951) if they still exist. Specimens of all the relatively common species {P. ^randis Rambur., P. pallida Glirrin, P. mai-ginata (Pawzlr), P. burmeisieriam Claassln) have been incorrectly determined as P. bipunctata. so the literature is hopelessly entangled. The Belgium specimen we list h as an egg that is generally similar to the egg of speci incns from Ireland and to a cariukiana KlAP-ALiai paraiectotype from Scotland, however, details of chorionic rcticulation suggest fivo species may be involved. A^ careful study of more material is needed and the possible selection of a neotype or use of plenary powers of the ICZN may be required to assure stability of this name. StDTOLLA - 2«: Figs, 1-3: SRM phototniciagTBphs of eggs; Peria bipuiiclMa Pictet [Ireland]: Egg lateral. 220x (J); iiüälarerd. 6W)x (2): detail of chorion and tnicropyles, $00\ P). "nir SfKCkcsofF.vickliv? tnn^ Ejfl Mtnplt^lng^' Figs. 4-6; SEM photomicrographs of eggs; Ferla hipunciom Pinn- [Balginni]: Egjj lateral, 24ÖX (4); eüMsr end, JOOx (5); delail of ehüriun and niicropyles, lOlOs {(i), SCOPOL1A49 -1002 The Peria burmeisteriana Group Ferlo. burmeisteriana Claasskn, 1936 Figs. 7-15 Egg.- Length 0,44-0.48 mm, width 0,28-0.30 mm, Collar short, about 0,02-0,03 mm long and (ibout 0.10-0,12 miti wide. Circumference of collar witli about 12 irregular thick ribs extending from rim to slioulderof egg body, ribs wide at rim aiid usually with a pit centered in the wide area at the rim: rim otherwise relatively smooth and only slightly Hanged. Chorion distinctly tii bereu late with prominent knob shaped processes covering entire surface; tuberculae vary in shape from rounded to irregularly linear outlines and the surface from smooth to emai^inate or deeply invaginatcd; an irregular lattice of low connectives and irregular pits surround tubcrculae. Micropylar orifices with thin raised rims set in low areas between tuberculae: micropylar row set about 0.16-0.20 mm from pole. SEM Specimen Data.- Austria, no specific locality (NMP). Bulgaria, Beli Viti, 16 May 1969, D. Braasch (SMNH). Czech Republic, Librice, 28 May !916, Komarek (NMP). Macedonia, Treska, Matka, 19 May 1976, P. Ikonomov (NMS). Montenegro, Ibar River, Rožaj, 26 May 1979,1. Sivec fSMNH). Switzerland, Plateau Vaudois, Broie Payeme, 15-21 May 1942, J. Aubert (MZ). Distribvuion.- Reported from Spain and North Africa i^orth to Holland and Luxemburg, east to the Carpathian Mountains of Ukraine and south to Macedonia. We have confirmed specimens from Austria, Bulgaria, Czech Republic, flermany, Hungary, Macedonia, Moiitene^ro, Slovenia and Switzerland. Comtnents,- This species was earlier recognised as P. abdominalis Burmeist^r before that name was found to be a junior homonym of P. ahchinimlis Guhrin and renamed by Claassen 0936). The holPt>'pe male from Germany is in the Museum Halle and specimens we have seen from Bavaria agree closely in egg morphology with Swiss and Austrian specimens. The remarkable egg of this species has been illustrated by Braastm & Joost (1971), Zs^'ICk (l 971) and Sivec et al. (1988). JH'j oC^rt,, (PlMpglna: J\j|iJii.); EtidcnL; fwm Eg^ Mmiilwliw Figs. 7-9: S£M phcuomicmgraphs of eggs: Perh (mrmei/:ieritma Claa-SSEN [SwiuerlondJ; Egg laleraU 2Ö0:( O)- collar end, 71 Ox (g); detai S of chorior e nd niicropy le, 131 Ov {9), SCOPOQA 19 . HKC Figs, 10-12: SEM pholomicrographs of eggs: Perla hummsleriam Cu^ASStN [Auslria]: F.gg lalcral, 24Qx (in); collar end. 5S0Jt (31); dctdil of chorion and iiiicropyles, Iö^Ok (12), •| kSpetil.H)fftTlljf(Ptm:in.:ni; Perlidac;: EviJeifci; Irtml kloqiimiagy KiKS. 13-15: SEM pliülomiciugraphs ofeg^s: PerSa huriK^hieriä'm CLAASiElv [Macedolliii]: R^g lateral, 24H>l (13); collav end, 55{)x (14); Jelail of chorion ajid rtitCfopi>ks, lOKQx (15). scopolja 49 - 2(102 Perla sp. nov. Sivec & Gr.\f* Ftgs. 16-IS Egg,- Lenglh 0.51-0.54 mm, width 0.2S-0.30 mm. Collar short, about 0.02-0,03 mm long and about 0.! 0-0.12 mm wide. Circumference of collar with about 12 irregular ihick ribs extending from rim and continuous with follicle cell iinpressions of egg body. Chorion covered tliroughout with a coarse lattice of raised follicle cell impression walls surrounding a finely pitted floor; cell innprcssion shape irregularly hexagonal and size varies from about 0.017 to about 0.029 mm in inner diaineter. Micropyles set in follicle cell impressions about 0.17-0.24 mm from pole; orifices with thin raised rims, SEM Specimen Data,- Austria, Kärnten Wölfnitz (SMNH). Austria, Vienna. 8 May 1999 (SMNH), Austria, Vienna, 17 May 1995 (SMNH). Slovenia, Ljubljana, 19 May 1899 (NMP). Distribution.- Kjiowu trom a few sites in Slovenia and Austria (Sivbc & Graf pers.com,), Comments.- This species is another in the P. burmeisieriana complex as Klapalek's detenmnation oiTI abdominalis for the Ljubljana specimen reflects. The egg collar and the chorionic surface perforations in the follicle cell impressions arc both suggestive of this association but the two are clearly distinct. The egg of this species is also somewhat similar to that of P hnrvati, described below from Turkish specimens, but iu that species only the coarser surface pits occur, * Detail description of species will be published in other paper. Tht Speti« ntf^rh msSi^cvt. PlsriHtael; Evidcntr from Eee Mi;rptnilu|;> Figs, 1613: SEM pholümitrogniphs of eggs; Fcr!a sp, nov. S]VEt:&ORA>- [Slovcnij)]; ilgg lalera!, :20k (Ift)i collar end. iSÜx (17); detail of chnriun and micropylcs, tÜSOx (IS). Perla zmckU sp. nov. Figs. 19-21 Types.- holotype, male: Turkey, Anatolia, I km NW Sogukptnar, lOOO m, 4.6.1992, leg. Malicky. Paratypes; 5 males, 3 females, same dala (Type material is ^lepo^iled in Slovenian Museum of Natural History, Ljubljana, Slovenia). Male.- Length of fore wings 19-21 iTim Fcinale.- Length of fore wings 25-29 mm Lgg,- Length 0.43-Ü.49 mm, width 0,2S-0.32 mm. Collar short, about 0.01 mm long and about 0.10-0.11 mm wide. Circumference of collar with about 12-16 modified follicle cell iitipressiüTis between rim ünd egg body. Chorion distinctly tubercukte with prominent, irrcgnlarly Sihiiped hexagonal plates covering entire surface; tuberciilae vary in size but those around equator are about 0.02-0,03 mm wide acros.? Ihe greatest distance; equator with about 12 plates in lateral aspect; plates rather tightly packed but surface surrounding plates with numerous shailow pits; some plates (about 30%) also have a single surface pit. Micropylar orifices with thin raised rim.s set in low areas between plates; micrapylar row set about 0.14-0.17 mm from pule. Etymology.- The patronym honors our friend and colleague Peter Zwick who, in ] 971. ha J already recognized and illustrated (in Fjg, 1 Od) this distinctive egg and speculated about the potential value of egg data in recognition of Peria species. Comments.- Although this species was placed as P. iliwsi Bka.^si; h & Joüst in Zwioi (1971) the egg is clearly more similar to burmeisteriana and we regard them as sister species. SCOI"nLW4; (2S); collar end. 550x (29); delait uf churiun. 1 ISÜx [30). The Perla gramJis Group Perlagrandis Ramblr, 1842 Figs. 31-33 Egg.- Length ü.42-0.47 min, width 0.25-0,30 mm. Collar short, about 0,02 mm long and about 0.09 mm wide, Circunifcrmcc of collar with about 24 irregularly spaccd vertical ribs extending from rijn onto shoulders of egg body and forming irregularly shafnjd, siomewhat trapezoidal cells an,iund shoulders. Rim very slightly Hanged and subtly incised, but these visible only in polar aspect. Chorion texture granular in appearance with fme detail showing irregular shon vermiform surface processes and very fine surface pitting over entire surface. Micropyles with inoderately large f'unneMikc orifiees surrounded by smooth low rims; micropylarrow set about 0.15-010 mm from pole, SEM Specimen Data - Austria, no other data (NMP). Poland, Zakopane, 14 July 1985, S.W. Szczytko (SWS). Slovenia, Lepena, ßovec, 2,5 May 1983,1, Sivec (SMNH). Switzerland, Valai, Col de Bretolet. August 1%3 (MZ). Distribution,- Reposled from the Sierra Nevada of Spam to Belgium and southeastern Poland in the north, through the Italian Peninsula to Sicily and as far cast as the Carpathian Mountains in Ukraine and Roumania and south as far as Croatia, We have confirmed specimens from Austria, Czech Republic. Italy, Poland, Serbia. Slovakia, Slovenia. Spain, Morocco and Switzerland. Comments.- Earl icr authors such as KlapAlek. (1923). Despax (1951), III !i:s(l955)andAuDiiRT (1959) rather consistently used the name Perla maxima (Scotoli) for this spccies, however, P. mcvcimn, through action of the ICZN "relative prcccdcni:e"pi"ocedure has been placed as a synonym of/? mij-gwata (Panzf.k). Conseglio (1967a) first proposed the suppression off! maxima and also applied the name, P. gramih to this species in Consigljo (1967b). StXiKJLlÄ w -axu JI Figs. 31-33: SEM pholomicrographs of eggs: FeHtt grandis RAitaufc [SwiLserkniil; Egs lüWra!, ZlO^ (3 H; collar enit SüOx (32); dew 11 of chorion and mkrapyles, 1[)4(!!( (33). Thü Speti» pr/Wj tPknjplmi: Fnliiljth; Eviiitn« from Ik Mwiiltilliny The PeHa horvati Group Per la horvati, sp, nov. Figs. 34-36 Types.- holotype, male; Turkey, r. Aksu, Tamdere, K urn bet, Se h it I er Gee, 1550 m, 2627,6,1995, kg. ITorvnt, Paratypes: 33 inales, 42 females, same data (Type material is deposited in Slovenian Muscuiti of Natural IJistory, Ljubljana, Slovenia). Male." Length of fore wings 22 mm Fe ma le,- Length of fore wings 28 mm Egg.- Length 0.59 mm, width Ü.35 mm. Collar short, about 0.02-0,03 mrn loi^g and 0.17 mm wide, Circumfercnce of collar without discemable ribs or follitie cell impressions; rim smooth and without flange or incisions. Chorion coarsely pitted; pUs iTTegularly oval or round in shape, relatively large (ca. Ü.017-0.029 mm diameter) and deep; surface between pits smooth; equatorial scction with about 10-12 pit rtjws in lateral aspect. Micropyles set in pits about 0.249 mm from pole; orifices .slightly raised with thin rims. Etymology." The patronym hontire our friend and colleague, B. i lorvat, curator at the Slovenian Museum of Natural History, and collector of the type series. SCOPOLIA 49 - MO: Fi^s. SEM pholumicrogyuphs of uggs: Perla harvali ip. nuv. Sivel&St/iRK flurkey): Egg lateral. 170x (34); collar erd, 3SOs (35J; detail of chorion aiid mioropyles, JtUx The Per la marginatu Group Perla marginaia (Panzer, 1799) Figs. 37-39 Egg.- Length 0.41 -0.46 mm, width (3.29-0,30 nun. Egg collar hardly produced, essentially a sessile opening surrouritJed by smooth tenninLs; basal width of tenninus about 0.06-0.08 mm. Chorion covered throughout with prominent to Hide cell impressions; hexagonal cells fairly uniform but cell widths ranging; frum 0,018-0.023 mm; cells lightly packed but separated from adjacent ones by narrow, ."ihallow sulci; cell floors with a central depression containing usually a single m in me pore; depressions house a single globular munhroom body whose stalk attaches to cbonon through oell pore. Micropylar row set about 0,13-0.16 mrn from pole; orifices without rims, set along intracellular borders at irregular points. SEM Specimen Data.- Germany, Eder Bergland n. Marburg. 9 June 1990, P. Zwick (LPS). Germany, Hammersbach, 5 June 1988, P. Zwick (LFS). Germany, Har^, Kinier Eberssberg, 12 June 1969, Tobias (LFS). Slovenia, Divje jezero, 8 June 1979, J, Sivec (SMNH). Slovenija, Mala Voda, Sevnik, 24 May 1990. B. Horvat (SMNH). Slovenia. Slahia. Grgar, 19 May 1993,1. Sivec (SMNH). Distribution.- Known from western Europe ranging from Spain to Holland east to Italy and western Poland. Earlier records from North Africa, Iran and the southern Balkans apparently pertain to P. pallida. We have confirmed specimens from Austria, Czech Republic, Germany, Italy, Poland, Slovakia, Slovenia and Spain. Comments." Eggs of this species are recognized on the basis of the extremely reduced col lar. In other respects the eggs are quite similar to those of P. matintemix Ramhur and to the P. palUda complex. TheSpcciea af fjrJa [?lgi:ap[Hi: [■|,nlidae>; F.vijcilK ticill Effl ManaialDgy Figs, 37-39; StM pholumitrographs of ejjgs P^^' la mcii^mula (Pamper) [Slovenial: Egg lateral, 2'IOx (37): collar end, ;2(lx t3R>; (Jelail of chorion and riicropy[cs, lOOOx (39), SCCIPÜLlAJf ■:«(! Perla madriten^is Rambur, 1842 Figs. 40-42 Egg.- Length [).42-0.45 mm, width 0,29-0.3(J tiim. Collar short, about 0,04 mm long, not distinctly offset from egg body and narrowed from basal width of about 0.13 mm to rim widtli of about 0.Ü9 mm; rim relatively smooth but, at least in some specimens, with indistinct folhcle cell impressions continuous from rim to egg body. Chorion covered throughout with prominent follicle cell impressions; hexagonal cells fairly unifomn but with cell widths ranging from 0.016-0.020 mm; ceils separated from adjacent ones by narrow sulci, and ceil floors containing a central depression and one or two minute pores, each cell depress i oil houses a single globular mushroom body whose staik attaches to the chorion through the ccH pores. Micropylar row set about U.l 5 -0.17 mm from pole; orifices without rims, set along intracellular borders at irregular points. SEM Specimen Data.- Portugal, Terras de Pontesa da Figulira, 3 June 1972 (LFS). Spain^ Gosende, 6 June 19S2, A. Conma (LFS). Distribution.- Reported from the basins of the Tego and Douro Rivers (Berti ielemy & Terra 1980) in ihe northern half of the Iberian Peninsula. We hstve confirmed spec i metis from Portugal and Spiim, Comments,- KlapAlek (1923) evidently had the holotype of this species from the "vicmity of Madrid" in his posession at one time but we have not located the spec mien and its existence has apparently not been documented since that time, for much of the last century the species was regarded as a synonyin ofmw-^inata but Aubert (1963) recognized it as a subspecies of P. mm-ginala and Berti eeltmy & Terra (195^0) considered it a valid species at least paitiaJly on the basis of egg morphology. More recently, MtMuiLLA (1990) suppoited this position by showing the drumming signals of P. madritensiii arc distinct from those of fl mar^imta P<:FlidBg]: Evtdtncc frcjr E^ MorpTifllogy Figs, 40-42: SEM photoiTiicro^raplis of Perh mtidnt^nsis Ramhlk [Spain]: Egg talcra], 240X (40); collar tniJ, (VOOx (41 J; detail (if chorion ard micTopyle, 1220k (42), Perla pallida Gvfm^, 1838 Figs. 43^63 Egg." Length 0.43-0.47 mm, width 0,27-0.30. Collar short, about Ü.02-0.03 mm long and about 0.09-0,12 mm wide; collarfonii variable but often, but notaJways, with the rim flanged and distinctly wider across the riin than the neck; sides of collar with ornamentation varying fi-om irregularrowsof pits to regular follicle cell impressions. Chorion covered throughontwitli prominent fo 11 i c I e eel I i mp rcssion s; hcxagona I eel I s fa i i l y u ni I'orm bi.i i ce 11 w i d ihs rangi n g froin about 0,0150.024 mill; cells typically separated from adjacent ones by narrow sulci hut in some specimens these arc rcduccd to a series of pits leaving incumplete sulci for cell impressions near collar; cell floors contain a ccntral depression and one or two minute pores; each cell depression houses usually one. but up to three mushroom bodies. Micropylarrow set about C. 15-0,22 mm from pole; orifices without rims, set along intracellular borders at irregular points, SEM Specimen Data - Type 1 Egg - Armenia, Sevan Basin. Argicki, 13 June I9fil {ZfSP), Caucasus, A. Kownackj, no additional data (LPS). Turkey, Macka, Trabzon, B, Hor^al (SMNH). Turkey, Kasianoan, Azdanaj, 21 June 1996, B, }Iorvat (SMNH), Turkey, Anatolia, no additional data (LPS), Turkey, Anatolia, Anat But Umg., Savi Goi, 7 August 1978 (LPS), Type 2 E^s -Bulgaria, 29 July 1969, D. Braasch (BPS). Bulgaria, Pirin, 3 June 1964, D. Braasch. Macedonia. TM. Stirevo Voda, 15 April 1999, B. Horvat (SMNH). Macedonia, Pesočani, II June 1979, P. Ikonomov (NMS), Macedonia, 20 July 1977, t. Sivec (SMMH). Slovenia, Gabrje, Žerovmk, 28 June 199S, i. Sivcc(SMNH}. Slovenia, Bisniea, nad Selnico, 6 June 1983 (SMNH), Type 3 Egs--Bulgaria, Stranea Gebirge. 15-19 June 1980, H. Malicky {LPS). Greece, Mega Peristeri, 22 May SLXJPOLEA 4<) - 2<10J 198S, J. Sivec, S. Lfchida (SMH N). Grcece, RiverTliiamis, Soulopoula, 21 May 1988,1. Sivec, S. Uchida (SMNH). Montenegro, Durmitor Moumain, 15 June 1983, C KruSnik (SMNH). Morocco, Moyen Atlas, del'Oued, Oum River, 4 May 1 %0, J. Aubert (MZ). Type4 Egg - Greece, Klepa, 25 June 1990. [. Sivec (SMNH), Distribution.- This species complex is reported from central Burope to the Caucasus, We have con firmed specimens from Armenia, Austria, Bosnia, Bulgaria, the Caucasus, Greece^ Macedonia. Montenefyro, Morocco, Slovakia, Slovenia and Turkey. Comments.- Perla paHida may be a complex of three or more species or subspecies. The original type ofpallida (now lost) was from the Caucasus and all egg samples we obsened fi-om the Caucasus. Turkey and Armenia had very similar egg collars oi" Type 1, with the rim flanged, incised and distinctly wider than the neck. Sides of these collar?; have prominent, continuous ribs extending from the shouldere to the rim. Type 2 egg collars occur on all samples examined from Slovenia and Macedonia and some from Bulgaria. These eggs have the collar about the same diameter (or slightly narrower) at the rim than the base so there is no constricted neck. Irregular thick nbs extend from the cell impi-ession mesh work of the egg body to the rim and these are defined by a series of conspicuous pits arranged in two or three irrcgulai' tiers, Type 3 collars are slightly shorter than those of Type 1 or 2 and there is a very slight constriction in the neck region and a slight expansion of the rim. Meshwork on the sides of the collar consists of narrow poorly defined ribs separated by a low suleus. Type 3 collars occur on specimens from Greece, Montenegro, Morocco, Bulgaria and from two samples labeled only "Yugoslavia". One additional egg sample which we tentatively place in this complex appears much more distinct and is referred to as Type 4, Cell impressions above and below the equatorial micropyles of this egg are outlined by a series of minute pores and faiitt lines which develop into a series of parallel lines of pits near the collar. This egg sample comes from two females captured in central Greece. Our egg data suggests P. paUuia is a complex in need of comprehensive review. A laiger sample of material inclusive of the entire range is needed and careful study of types of two current synonyms R dacica Ki.ap.^let and burexchi Schoenemlnd must be included. nn SjWClBi wr ATVg PL-rtiifae); Evidiaici: Tmrn Egg Miin;1ialti8}- Figs. 43-4S; SEM phOtomieroersphsofeBg-':: PcilapaiUdu GutJils, Type 1 [Armenia); Egg latwal, iJOs (43), collar end. STOjt detail of chorion ard micnapylcs, lOOOx (45). scofölla ■i? - Figs. 4(1-«: SEM phowniicmgrarihs of efigs: PerlapatMa Gleiün. Type 1 [Turkey]: Egg lateral, 5,10s (46): tollar end, 600x (47); desiil of cliorion imd micmpyle, 1210k (4Ä), "nie Spocies of AWj (PtetMptera: Pertidacj: Evidtinfc ftptn Egg MprpfagloEy Figs. 49-51: SEM pholumitrügraphs of eggs; P^'^/^I/J^?//^^/[■lOuERl^, Type 2 (Bulgaria)' Egg laleral, 240x collar erd 500x (iO); dc[Bil ort-horion and tnkrapyles, lÜlOx (51). SnjpOUAW-IlKß Figs. 52-54: SBM pholomteogn^hs of c^s: t'ertii pallida üutdi^, Type 2 [Macedoni j]: Egg lateral, lAOr. (52); i;oltar end, SOOx [S3); detail of choricn, 12]Ox (45), Tlic SfflCia iifA'WM iFlKaipteni: Piriidati: EviilfiKi: Ikmi EffiMpfTholpgy 0'SA Figs !5-S7: SEM phutomicmgraplis of e^: PedapaUisja Curaix,Type2 [Slovenia]; Egg lateral. 240x (55); tollarend, 470jc (56); detail of chorion arid micropyle, 121 Ox (57), SrOPOLlA - 2003 Xf r ' i fC 60 Figs, 58-60: SEM photomitnugraplis ofe^gs; FerlapaliiiSa Gufrin, Type 3 [GrecccJ: Egg lalerat, 2-(0x (5S); eoJliU and, 630s (59); detail of chorion and niicropylei. tOlOx (60). The Sjtcdts ff rill Ifli.'Miplna: Pl;rlijBf)i: EviifcnLe fimr Mmjiillllllgy Figs. 61-Ö3: SEM photoraicrograpbs of eggs: Peila puiUJa Oufrin. Type 4 [Oreecci: Ej^g laltral. 220s (61); collar end Sl>0\ (63); cltfail of chorion and riicmpylas, 121 Ox Povzetek Predsiavniki roduP^j /o sodijo med najveCje evropske vrbnice, Perla i^ najstarejše rodovno ime med vrbnicaini in pod tem imenom so bile opisane številne vrste. V katalogu Illjesa (1966) za ta rod zasledimo kar 293 imen vrst, ki večinomn sodijo jncd vprašljive vrste. Sevec in ostali (1988) so rod Psrla omejili le na Evropo ii: nekatere sosednje dežele, tako da se je Število vrst znotraj rodu znižajo na 8. Študij precej obsežnega materiala v preteklem desetletju je pokazal, da je taksonomija tega redu se vedno zelo nejasna. V tem delu pred stav Ijame sistematski pregled posameznih vrst rodu Perla no podlagi Študije strukture površine jajc posameznih vrst z elektron.skiiii rastrskim naikroskoponv Ugotovili smo 12 vrst, ki jih uvrščanto v šest skupin vrst. Perla korvati in Perla zwicki iz TtirČije sta za znanost novi vrsti. 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