Original scientific article UDK 576.89:593.1 Received: 2013-03-26 NOTES ON THE GREGARINES (PROTOZOA: APICOMPLEXA: EUGREGARINORIDA) OF INSECTS IN SLOVENIA Dušan DEVETAK Department of Biology, Faculty of Natural Sciences and Mathematics, University of Maribor, SI-2000 Maribor, Koroška cesta 160, Slovenia E-mail: dusan.devetak@guest.arnes.si Manja OMERZU SI-2000 Maribor, Borova vas 29, Slovenia Richard E. CLOPTON Department of Natural Science, Peru State College, Peru, Nebraska 68421, U.S.A. ABSTRACT Gregarines (Apicomplexa: Eugregarinorida) are relatively large protozoan parasites of the guts and body cavities of invertebrates including annelids, tunicates, sipunculids, and especially arthropods. The knowledge of their occurrence in insects is poor; gregarines have been reported from less than one percent of named insect species. Between August 2011 and October 2012 intestine of larvae and adults of insects originating from Slovenia were eviscerated and inspected for gregarines. Hosts of the following orders were infected: Dermaptera, Orthoptera, Blattaria, Psocoptera, Neuroptera and Coleoptera. During this preliminary study, 20 gregarine species were recorded belonging to the following taxa: Actinocephalus, Euspora, Gamocystis, Gregarina, Hirmocystis, Hyalospora and Leidyana. Key words: Gregarines, Eugregarinorida, Apicomplexa, insects, parasites, Slovenia NOTE SU GREGARINE (PROTOZOA: APICOMPLEXA: EUGREGARINORIDA) DI INSETTI IN SLOVENIA SINTESI Le gregarine (Apicomplexa: Eugregarinorida) sono protozoi relativamente grandi e parassiti del tratto digerente e delle cavita corporee di vari invertebrati, inclusi anellidi, tunicati, sipunculidi e specialmente artropodi. Molto poco e finora noto riguardo alla loro presenza negli insetti, visto che le gregarine sono state ritrovate in meno dell'un percento delle specie di insetti conosciute. La ricerca si e svolta nel periodo fra l'agosto del 2011 e l'ottobre del 2012, quando sono stati eviscerati ed ispezionati, alla ricerca di gregarine, i tratti digerenti di larve ed adulti di insetti provenienti dalla Slovenia. Individui dei seguenti ordini sono stati infettati: Dermaptera, Orthopte-ra, Blattaria, Psocoptera, Neuroptera e Coleoptera. Durante questo studio preliminare sono state riconosciute 20 specie di gregarine, appartenenti ai seguenti taxa: Actinocephalus, Euspora, Gamocystis, Gregarina, Hirmocystis, Hyalospora e Leidyana. Parole chiave: Gregarine, Eugregarinorida, Apicomplexa, insetti, parassiti, Slovenia INTRODUCTION Gregarines (Protozoa: Apicomplexa) are obligate unicellular parasites infecting the intestines and other organs of invertebrates of terrestrial, freshwater, and marine habitats. The majority of eugregarine species are reported from insects (Clopton, 2002; Rueckert & Leander, 2008). The knowledge of their occurrence in insects is poor and is currently not given the attention they deserve. Therefore, most gregarine species remain unknown and undescribed. Gregarines are divided into three major groups: eu-gregarines, archigregarines, and neogregarines (Grassé, 1953; Levine, 1971; Leander, 2008). Despite recent studies on the molecular level (see Clopton, 2009), our understanding and delineation of supra-specific taxa is incomplete because of our poor understanding of the actual diversity and phylogenetic relationships of grega-rines (Rueckert & Leander, 2009). Although taxonomic keys, figure drawings, or micrographs of many European gregarine species are given in the monographs of Lipa (1967) and Geus (1969), our knowledge of the diversity and distribution of the European gregarine fauna is still poor. In this study, gregarines from insects in Slovenia are addressed for the first time. MATERIAL AND METHODS Adult and larval insects collected in the area of Maribor (NE Slovenia) were eviscerated and their alimentary canals dissected in insect saline. Individuals of the following insect orders were inspected: Dermaptera, Orthoptera, Blattaria, Psocoptera, Neuroptera and Coleoptera. Their intestine was examined microscopically at 100, 200 and 400-times magnification. Gregari-nes were measured and photographed using a Nikon E 800 Microscope with a mounted digital camera Nikon DN100, and Eclipse Net version 1.16.3 software. The following standard gregarine trophozoite and gamont metrics (Lipa, 1967; Clopton,2004) are herein reported (in |jm): total length, length of epimerite, length of proto-merite, length of deuromerite, maximum width of pro-tomerite, width of deutomerite at equatorial axis, and maximum width of deutomerite. RESULTS AND DISCUSSION One hundred twenty-two individual hosts' taxa within 6 insect orders were examined and twenty gregarine species were recorded and addressed herein. Order Eugregarinorida Léger, 1900 Superfamily Gregarinoidea Chakaravarty, 1960 Emend. Clopton, 2009 Family Gregarinidae Labbé, 1899 Genus Gregarina DufouR, 1828 (Plates I, II) Gregarina ovata Dufour, 1828 (Figs. 1, 2) Material examined: Host: Forfícula auricularia (Linnaeus, 1758) (Dermaptera: Forficulidae) Locality and prevalence: Maribor, 30.8.2011, 2/2 individuals infected; 9. 9.2011, 4/8 individuals infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are presented in Table 1. Clopton et al. (2008) reviewed morphometric data and nomenclatural status of previously described species of Gregarina infecting earwigs and recognized 8 valid species, two of which are reported from Slovenia. G. ovata was also reported from F. auricularia by Geus (1969). Locality reports (summarized in Geus, 1969; Clopton et al., 2008): Africa (Cabo Verde Islands), Asia (Japan), Europe (France, Germany, Great Britain, Poland) and North America (USA). Gregarina chelidurellae Geus, 1969 (Fig. 3) Species inquirenda Material examined: Host: Forfícula auricularia (Linnaeus, 1758) (Dermaptera: Forficulidae) Locality and prevalence: Maribor, 9. 9.2011, 1/4 individuals infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. G. chelidurellae is differentiated from G. ovata by differences in gamont size and shape. Geus (1969) provided only morphometric data and description for gamonts and associations but no oocyst description or data thus the taxon is considered species inquirenda (Clopton et al., 2008). G. chelidurellae was also reported from Chelidura (=Chelidurella) acanthopygia (Dermaptera) by Geus (1969). Locality reports (summarized in Geus, 1969; Clopton et al., 2008): Europe (Germany). Gregarina acridiorum (Léger, 1893) (Fig. 4) Material examined: Host: Decticus verrucivorus (Linnaeus, 1758) (Orthoptera: Tettigoniidae) Locality and prevalence: Maribor, 16.10.2012, 2/4 individuals infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. G. acridiorum is a common species occurring in locusts and grasshoppers (Orthoptera) (Geus, 1969; Lipa et al., 1996). Locality reports (summarized in Geus, 1969; Lipa et al., 1996): Europe (France, Germany, Poland), Africa (Algeria) and Asia. PLATE I: Genus Gregarina Fig. 1: Solitary individual of G. ovata Fig. 2: Associative pair of G. ovata Fig. 3: Solitary individual of G. chelidurellae Fig. 4: Solitary individual of G. acridiorum Figs. 5, 6: G. delmasi; (Fig. 5) association and (Fig. 6) syzygy TABLA I: Rod Gregarina Sl. 1: Solitarni osebek vrste G. ovata Sl. 2: Ascociaciji vrste G. ovata Sl. 3: Solitarni osebek vrste G. chelidurellae Sl. 4: Solitarni osebek vrste G. acridiorum Sl. 5,6: G. delmasi; (Sl. 5) asociacija in (Sl. 6) sizigij Gregarina delmasiTuzet et Rambier, 1953 (Figs. 5-8) Material examined: Host: Decticus albifrons (Fabricius, 1775) (Orthopte-ra: Ensifera: Tettigoniidae) Locality and prevalence: Maribor, 5.9.2011, 17.10.2011, 2/2 individuals infected. Gametocyst diameter 244-260 pm (n=4) (Fig. 7). Ga-monts with characteristic longitudinally grooved pellicle (Fig. 8). Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. G. delmasi was reported in locusts and grasshoppers (Orthoptera) by Geus (1969). Locality reports (summarized in Geus, 1969): Africa (Congo) and Europe (France). Gregarina katherina Watson, 1915 (Fig. 9) Material examined: Host: Adalia bipunctata (Linnaeus, 1758) (Coleoptera: Coccinellidae) Locality and prevalence: Maribor: Piramida, 11.9.2011 2/2 individuals infected. Associations composed of 2-3 individuals, caud-ofrontal. Measurements of gamonts in associations are given in Table 1. G. katherina is reported from the following cocci-nelid species (Coleoptera: Coccinellidae) (Watson, 1915; Geus, 1969; Hoshide, 1980): Coccinella septempunctata, Coccinula quatuordecimpustulata, and Coccinella no-vemnotata. Locality reports (summarized in Lipa, 1967; Geus, 1969; Hoshide, 1980): Asia (Japan), Europe (Germany, Poland) and North America (USA). Gregarina barbarara Watson, 1915 (Fig. 10) Material examined: Host: Adalia bipunctata (Linnaeus, 1758) (Coleoptera: Coccinellidae) Locality and prevalence: Maribor, Piramida, 28.9.2011, 2/6 individuals infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. G. barbarara is reported from the following cocci-nellid species (Watson, 1915; Geus, 1969): Adalia bipunctata, Coccinella sp., Brumus quadripustulatus and Tyt-thaspis sedecimpunctata. Locality reports (summarized in Geus, 1969): Europe (Germany) and North America (USA). Gregarina steini Berndt, 1902 (Figs. 11, 12) Material examined: Host: Tenebrio molitor Linnaeus, 1758: larva (Coleoptera: Tenebrionidae) Locality and prevalence: Maribor, from April to May 2012; 9/20 larvae infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. Gregarines occurring in mealworm (T. molitor) larvae and adults are detailed by Clopton et al. (1991, 1992) and Clopton & Janovy (1993). G. steini is known only from T. molitor. Locality reports (summarized in Geus, 1969; Lipa, 1967): Europe and North America (USA). Gregarina cuneata Stein, 1848 (Figs. 13-15) Material examined: Host: Tenebrio molitor Linnaeus, 1758: larva (Coleoptera: Tenebrionidae) Locality and prevalence: Maribor, from April to May 2012; 4/20 larvae infected. Association biassociative, caudofrontal. Triassociati-ve associations occurred rarely (Fig. 14). Measurements of trophozoites and gamonts in association are given in Table 1. G. cuneata is reported from the following tenebri-onid beetles (Coleoptera: Tenebrionidae) (Geus, 1969): Alphitobius ovatus, Ceropria anthracina, Ceropria ro-mandi, Chiroscelis digitata, Stenosis angustata, Stron-gylium buettneri, T. molitor, Tenebrio nitidulus, Tenebrio obscurus and Tribolium ferrugineum. Locality reports (summarized in Geus, 1969; Lipa, 1967): Africa, Asia (Turkey, Japan), Europe, North and South America. Gregarina polymorpha (Hammerschmidt, 1838) (Fig. 16) Material examined: Host: Tenebrio molitor Linnaeus, 1758: larva (Coleoptera: Tenebrionidae) Locality and prevalence: Maribor, from April to May 2012; 14/20 larvae infected. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. G. polymorpha is reported from the following tene-brionid beetles (Coleoptera: Tenebrionidae) (Geus, 1969): Gonocephalum sp., Gonocnemis sp., Pelloides senega-lensis, Taraxides punctatus, T. molitor and Tenebrio gu-ineensis. Locality reports (summarized in Geus, 1969; Lipa, 1967): Africa (Congo), Asia (Japan), Europe, North and South America. PLATE II: Genus Gregarina Figs. 7,8: G. delmasi; (Fig. 7) gametocyst and (Fig. 8) detail of the pellicle of a primate Fig. 9: Association of G. katherina Fig. 10: Association of G.a barbarara Figs. 11,12: Solitary individual and association of G. steini Figs. 13-15: G. cuneata; (Fig. 13) biassociative association, (Fig. 14) triassociative association and (Fig. 15) solitary individual Fig. 16: Association of G. polymorpha TABLA II: Rod Gregarina SI. 7,8: G. delmasi; (SI. 7) gametocista in (SI. 8) detajl pelikule primita SI. 9: Asociacija vrste G. katherina SI. 10: Asociacija vrste G. barbarara SI. 11,12: SoIitarni osebek in asociacija vrste of G. ste- ini SI. 13-15: G. cuneata; (SI. 13) biasociativna asociacija, (SI. 14) triasociativna asociacija in (SI. 15) soIitarni osebek SI. 16: Asociacija vrste G. polymorpha Gregarina curvata (Hammerschmidt, 1838) (Fig. 17) Material examined: Host: Cetonia sp. (Coleoptera: Scarabaeidae) Locality and prevalence: Maribor, 14.10.2011, 1/6 individuals infected. Measurements of trophozoites are given in Table 1. Gregarina curvata is reported from the following scarabaeid beetles (Coleoptera: Scarabaeidae) (Geus, 1969): Cetonia sp., Cetonia aurata, Protaetia cupraea (= Potosia cuprea) and Osmoderma eremita. Locality reports (summarized in Geus, 1969): Europe (France, Germany). Genus Gamocystis Schneider, 1875 (Plate III) Gamocystis tenax Schneider, 1875 (Fig. 18) Material examined: Host: Ectobius lapponicus (Linnaeus, 1758) (Blattaria: Blatellidae) Locality and prevalence: Maribor, 5.9.2011, 19.9.2011; 4/8 individuals infected. Association biassociative, caudofrontal. Only in younger trophozoites was the gregarine divided into a distinct protomerite and deutomerite. Measurements of trophozoites and gamonts in association are given in Table 1. Gregarines occurring in tropical and subtropical Blattaria are comprehensively presented in a series of papers of Clopton (1995, 2010, 2011, 2012a, 2012b), Clop-ton & Gold (1996) and Clopton & Hays (2006). Little is known about gregarines infecting Holarctic cockroaches. G. tenax is reported from the following cockroach species (Geus, 1969): Ectobius lapponicus, Ectobius livens and Ectobius sylvestris. Locality reports (summarized in Geus, 1969): Europe (France, Germany). Gamocystis fimetarii Cordua, 1953 (Fig. 19) Material examined: Host: Ectobius lapponicus (Linnaeus, 1758) (Blattaria: Blatellidae) Locality and prevalence: Maribor, 27.9.2011; 2/4 individuals infected. Association biassociative. The primite is clearly divided into protomerite and deutomerite, however, there is no such division of the satellite. Measurements of trophozoites and gamonts in association are given in Table 1. G. fimetarii was originally reported from an aphodiid beetle Aphodius fimetarius (Coleoptera: Aphodiidae) (Geus, 1969), so it is unusual that it is found in cockroaches. Locality reports (summarized in Geus, 1969): Europe (Germany). Family Hirmocystidae Grasse, 1953 Genus Hirmocystis Labbe, 1899 (Plate III) Hirmocystis polymorpha (Leger, 1892) (Figs. 20, 21) Material examined: Host: Adalia bipunctata (Linnaeus, 1758) (Coleoptera: Coccinellidae) Locality and prevalence: Maribor: Piramida, 12.9.2011; 2/4 individuals infected. Associations caudofrontal, composed of 6-7 individuals. Measurements of gamonts in associations are given in Table 1. H. polymorpha is reported from larvae of a lucanid beetle Platycerus caraboides (Coleoptera: Lucanidae) and from larvae of dipterans Erioptera sp., Limonia sp. and Symplecta sp. (Geus, 1969). This is the first report of H. polymorpha from a coccinellid host. Locality reports (summarized in Geus, 1969): Europe (Germany). Genus Hyalospora Schneider, 1875 (Plate III) Hyalospora hemerobii Geus, 1969 (Figs. 22, 23) Material examined: Host: Hemerobius humulinus Linnaeus, 1758 (Neu-roptera: Hemerobiidae) Locality and prevalence: Maribor, Piramida; 8.9.2011, 16.9.2011; 9/10 individuals infected. Gametocyst diameter 83-95 pm (n=4) (Fig. 23). Some trophozoites with epimerite. Association biassociative, caudofrontal. Measurements of trophozoites and gamonts in association are given in Table 1. H. hemerobii is also reported from the brown la-cewing species Hemerobius pini (Neuroptera: Hemerobiidae) (Geus, 1969). Locality reports (summarized in Geus, 1969): Europe (Germany). Hyalospora psocorum (Siebold, 1839) (Fig. 24) Material examined: Host: Graphopsocus cruciatus (Linnaeus, 1768) (Pso-coptera: Stenopsocidae) Locality and prevalence: Maribor: Piramida; 14.10.2012, 23.10.2012; 7/12 individuals infected. Measurements of trophozoites are given in Table 1. H. psocorum is reported from the following psocop-teran species: Amphigerontia bifasciata, Graphopsocus cruciatus, Mesopsocus unipunctatus, Psocus longicor-nis, Stenopsocus immaculatus, Lachesilla quercus and Valenzula flavidus (Lipa, 1967; Geus, 1969). Locality reports (summarized in Lipa, 1967; Geus, 1969): Germany (Poland). PLATE III: Genera Gregarina, Gamocystis, Hirmocystis and Hyalospora Fig. 17: Solitary individual of Gregarina curvata Fig. 18: Association of Gamocystis tenax Fig. 19: Associations of Gamocystis fimetarii Figs. 20,21: Association of Hirmocystis polymorpha Figs. 22, 23: Hyalospora hemerobii; (Fig. 22) solitary individuals and (Fig. 23) gametocyst Fig. 24: Solitary individual of Hyalospora psocorum TABLA III: Rodovi Gregarina, Gamocystis, Hirmocystis in Hyalospora Sl. 17: Solitarni osebek vrste Gregarina curvata Sl. 18: Asociacija vrste Gamocystis tenax Sl. 19: Asociacija vrste Gamocystis fimetarii Sl. 20,21: Asociacija vrste Hirmocystis polymorpha Sl. 22,23: Hyalospora hemerobii; (Sl. 22) solitarni osebki in (Sl. 23) gametocista Sl. 24: Solitarni osebek vrste Hyalospora psocorum Genus Euspora Schneider, 1875 (Plate IV) Euspora fallax Schneider, 1875 (Figs. 25, 26) Material examined: Host: Melolontha sp., larva (Coleoptera: Scarabae-idae) Locality and prevalence: Maribor, 3.5.2012; 2/4 individuals infected. Association biassociative, caudofrontal. Measurements of associations are given in Table 1. E. fallax is reported from the following beetles (Coleoptera) (Geus, 1969): Tenebrionidae: Allecula sp., Asida sp., Monomma giganteum; larval Melolonthidae: Melolontha melolontha, Melolontha sp., Rhizotrogus aestivus and Rhizotrogus sp. Locality reports (summarized in Geus, 1969): Europe (Germany, France) and Africa (Congo). Superfamily Stenophoroidae Levine, 1984 Emend. Clopton, 2009 Family Leidyanidae Kudo, 1954 Genus Leidyana Watson, 1915 (Plate IV) Leidyana gryllorum (Cuenot, 1897) = Leidyana erratica (Crawley, 1907) (Figs. 27, 28) Material examined: Host: Gryllus campestris Linnaeus, 1758 (Orthopte-ra: Gryllidae) Locality and prevalence: Kamnica, Koblarjev zaliv; 7.9.2011; 2/2 individuals infected. Gametocyst diameter 95-115 pm (n=6) (Fig. 28). Measurements of trophozoites are given in Table 1. L. gryllorum is reported from the following cricket species (Orthoptera: Gryllidae) (Geus, 1969): Acheta domesticus, Gryllus campestris, Gryllus assimilis, Ne-mobius fasciatus, Nemobius sylvestris, Phaeophilacris pilipennis and Phaeophilacris sp. Locality reports (summarized in Geus, 1969): Europe (France, Germany), USA, Asia (India) and Africa (Congo). Leidyana oblonga (Dufour, 1837) (Fig. 29) Material examined: Host: Acheta domesticus (Linnaeus, 1758) (Orthoptera: Gryllidae) Locality and prevalence: Maribor, a stock; 7.9.2011; 5/10 individuals infected. Measurements of trophozoites are given in Table 1. L. oblonga is reported from the following orthop-teran species (Orthoptera) (Geus, 1969): Gryllidae: G. campestris, N. sylvestris; Acrididae: Locusta migratoria, Psophus stridulus. Locality reports (summarized in Geus, 1969): Europe (France, Germany) and North America (USA). Family Actinocephalidae Léger, 1892 Subfamily Actinocephalinae Léger, 1899 Genus Actinocephalus Stein, 1848 (Plate IV) Actinocephalus permagnus Wellmer, 1911 (Figs. 30-32) Material examined: Host: Carabus coriaceus Linnaeus, 1758 (Coleoptera: Carabidae) Locality and prevalence: Maribor, 23.9.2011; 2/4 individuals infected. Protomerite cylindrical or conical (Figs. 31, 32). Measurements of trophozoites are given in Table 1. A. permagnus is reported from the following carabid species (Coleoptera: Carabidae) (Geus, 1969; Sienkiewi-cz & Lipa, 2009): Carabus cancellatus, C. catenulatus, C. coriaceus, C. granulatus, C. hortensis, C. nemoralis, Carabus (Cathoplius) asperatus and Ceutosphodrus oblongus. Locality reports (summarized in Geus, 1969; Sienki-ewicz & Lipa, 2009): Europe (France, Germany, Poland) and Africa (Morocco). Actinocephalus conicus (Dofour, 1837) (Figs. 33-35) Material examined: Host: Dorcus parallelipipedus (Linnaeus, 1785) (Coleoptera: Lucanidae) Locality and prevalence: Maribor, 10.6.2012, 31.8.2012; 1/4 individuals infected. Measurements of individuals are given in Table 1. Some trophozoites with epimerite. Actinocephalus conicus is reported from the following beetle (Coleoptera) species: Dorcus parallelipipedus (Lucanidae); Elater ferrugineus (larvae) (Elate-ridae). Locality reports (summarized in Geus, 1969): Europe (France, Germany). ACKNOWLEDGEMENTS We are grateful two anonymous referees for critical remarks. This project was supported by the Slovenian Research Agency within the Biodiversity Research Programme (Grant No. P1-0078). 27 • PLATE IV: Genera Euspora, Leidyana and Actinocepha-lus Figs. 25,26: Associations of E. fallax Figs. 27,28: L. gryllorum; (Fig. 27) solitary individual and (Fig. 28) gametocyst Fig. 29: Solitary individual of L. oblonga Figs. 30-32: A. permagnus; (Fig. 30) solitary individual and (Figs. 31,32) protomerites Figs. 33-35: A. conicus; (Fig. 33) individuals without epimerite and (Figs. 34, 35) individuals with epime-rite TABLA IV: Rodovi Euspora, Leidyana in Actinocephalus SI. 25,26: Asociacija vrste E. fallax SI. 27, 28: L. gryllorum; (SI. 27) solitani osebek in (SI. 28) gametocista SI. 29: Solitarni osebek vrste L. oblonga SI. 30-32: A. permagnus; (SI. 30) soIitarni osebek in (SI. 31,32) protomeriti SI. 33-35: A. conicus; (SI. 33) osebki brez epimerita in (SI. 34,35) osebki z epimeritom K POZNAVANJU GREGARIN (PROTOZOA: APICOMPLEXA: EUGREGARINORIDA) IZ ŽUŽELK V SLOVENIJI Dušan DEVETAK Oddelek za biologijo, Fakulteta za naravoslovje in matematiko, Univerza v Mariboru, SI-2000 Maribor, Koroška cesta 160 E-mail: dusan.devetak@guest.arnes.si Manja OMERZU SI-2000 Maribor, Borova vas 29 Richard E. CLOPTON Department of Natural Science, Peru State College, Peru, Nebraska 68421, ZDA POVZETEK Gregarine (Protozoa: Apicomplexa: Eugregarinorida) so razmeroma velike praživali, ki zajedajo v kopenskih, sladkovodnih in morskih nevretenčarjih. So obligatni paraziti v prebavilih in telesnih votlinah kolobarnikov, pla-ščarjev, pršivcev in členonožcev. Čeprav so iz žuželk opisali največ vrst gregarin, so za prisotnost teh parazitov preiskali le manj kot odstotek znanih vrst žuželk, zato pričakujemo še veliko novih najdb. Kljub bogati tradiciji v preteklosti so te praživali v Evropi še vedno razmeroma slabo preiskane. V prispevku so za Slovenijo gregarine prvič navedene. Med avgustom 2011 in oktobrom 2013 smo odkrili gregarine v prebavilih naslednjih redov žuželk: Dermaptera, Orthoptera, Blattaria, Psocoptera, Neuroptera in Coleoptera. Zabeležili smo dvajset vrst gregarin, ki jih uvrščamo v naslednje rodove: Actinocephalus, Euspora, Gamocystis, Gregarina, Hirmocystis, Hyalospora in Leidyana. 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Janovy Jr. (1993): Developmental niche structure in the gregarine assemblage parasitizing Tenebrio molitor. J. Parasitol., 79(5), 701-709. Clopton, R. E. & R. E. Gold (1996): Host specificity of Gregarina blattarum von Siebold, 1839 (Apicomplexa: Eugregarinida) among five species of domiciliary cockroaches. J. Invertebr. Pathol., 67, 219-223. Clopton, R. E. & J. J. Hays (2006): Revision of the genus Protomagalhaensia and description of Protomagalhaensia wolfi n. comb. (Apicomplexa: Eugregarinida: Hirmocystidae) and Leidyana haasi n. comb. (Apicomplexa: Eugregarinida: Leidyanidae) parasitizing the lobster cockroach, Nauphoeta cinerea (Dictyoptera: Blaberidae). Comp. Parasitol., 73(2), 137-156. Clopton, R. E., T. J. Percival & J. Janovy Jr. (1991): Gregarina niphandrodes n. sp. (Apicomplexa: Eugregarinorida) from adult Tenebrio molitor (L.) with oocyst descriptions of other gregarine parasites of the yellow mealworm. J. Protozool., 38(5), 472-479. Clopton, R. E., J. Janovy Jr. & T. J. Percival (1992): Host stadium specificity in the gregarine assemblage parasitizing Tenebrio molitor. J. Parasitol., 78(2), 334-337. Clopton, R. E., T. J. Cook & J. L. Cook (2008): Gregarina tropica n. sp. (Apicomplexa: Eugregarinorida: Gre-garinicae: Gregarinidae) parasitizing the brown-winged earwig, Vostox brunneipennis (Dermaptera: Labiidae), in the Texas Big Thicket. Comp. Parasitol., 75(2), 215-227. Geus, A. (1969): Sporentierchen, Sporozoa: die Gre-garinida der land- und süßwasserbewohnenden Arthropoden Mitteleuropas. In: Dahl, F. (ed.): Die Tierwelt Deutschlands und der angrezenden Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise. Gustav Fischer Verlag, Jena, pp. 1-608. Grassé, P.-P. (1953): Classe des grégarinomorphes (Gregarinomorpha, N. nov., Gregarinae Haeckel, 1866; gregarinidea Lankester, 1885; grégarines des auteurs). In: Grassé, P.-P. (ed.): Traité de Zoologie. Masson, Paris, pp. 590-690. Hoshide, K. (1980): Notes on the Gregarines in Japan 11. A new and two already-known species of Gregarines from Japanese Coccinellidae. Bull. Faculty Educ., Yama-guchi University, vol. 30(2), 49-60. Leander, B. S. (2008): Marine gregarines-evolutiona-ry prelude to the apicomplexan radiation? Trends Parasitol., 24, 60-67. Levine, N. D. (1971): Taxonomy of Archigregarinorida and Selenidiidae (Protozoa, Apicomplexa). J. Protozool., 18, 704-717. Lipa, J. J. (1967): Studies on gregarines (Gregarino-morpha) of arthropods in Poland. Acta Protozool., 5, 97-179. Lipa, J. J., P. Hernandez-Crespo & C. Santiago-Alvarez (1996): Gregarines (Eugregarinorida: Apicomplexa) in natural populations of Dociostaurus maroccanus, Cal-liptamus italicus and other Orthoptera. Acta Protozool., 35, 49-59. Rueckert, S. I. & B. S. Leander (2008): Gregarina. Gregarines. Version 23 September 2008. http://tolweb.org/ Gregarina/124806/2008.09.23 In: The Tree of Life Web Project, http://tolweb.org/ Rueckert, S. I. & B. S. Leander (2009): Molecular Phylogeny and Surface Morphology of Marine Archi-gregarines (Apicomplexa), Selenidium spp., Filipodium phascolosomae n. sp., and Platyproteum n. g. and comb. from North-Eastern Pacific Peanut Worms (Si-puncula). J. Eukaryot. Microbiol., 56(5), 428-439. Sienkiewicz, P. & J. J. Lipa (2009): Prevalence of eu-gregarines (Apicomplexa: Eugregarinida) parasitizing in ground beetles (Coleoptera, Carabidae) in various habitats. Pol. J. Entomol., 78, 351-368. Watson, M. E. (1915): Some new gregarine parasites from Arthropoda. J. Parasitol., 2, 27-36. Tab. 1: Measurements of gregarines. Abbreviations: TL - total length; EL - epimerite length; PL - length of protomerite; PL* - length of protomerite without epimerite; DL - length of deutomerite; PWM - maximum width of protomerite; DWE - width of deutomerite at equatorial axis; DWM - maximum width of deutomerite; Pr - primite; Sat - satellite; Assoc - association Tab. 1: Meritve gregarin. Okrajšave: TL - celotna dolžina; EL - dolžina epimerita; PL - dolžina protomerita; PL* - dolžina protomerita brez epimerita; DL - dolžina devtomerita; PWM - največja širina protomerita; DWE - širina devtomerita na ekvatorialni ravnini; DWM - največja širina devtomerita; Pr - primit; Sat - satelit; Assoc - asociacija Gregarina ovata Dufour, 1828 Solitary individuals Individual TL PL DL PWM DWE DWM 1 392 62 330 105 244 248 2 248 45 203 64 117 117 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 39 60 330 100 250 260 370 40 330 130 250 255 2 400 50 350 120 265 270 377 27 350 140 270 275 3 200 35 165 50 100 110 196 24 172 50 95 96 4 400 56 344 119 271 273 386 29 357 135 279 279 5 392 46 346 112 270 272 381 27 354 136 274 274 Gregarina chelidurellae Geus, 1969 Solitary individuals Individual TL PL DL PWM DWE DWM 1 127 26 101 32 53 53 2 201 35 166 44 71 72 3 156 34 122 35 62 62 4 172 36 136 35.5 66 66 5 252.5 42.5 210 61.5 98 100 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 212 38 174 41 62 63 228 36 192 53 69 69 2 211 38 173 42 65 65 227 37 190 52 69 70 Gregarina acridiorum (Léger, 1893) Solitary individuals Individual TL PL DL PWM DWE DWM 1 419 89 330 115 158 158 2 450 96 354 128 158 159 3 622 129 493 186 292 302 4 397 100 297 127 181 198 5 382 79 303 113 172 187 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 307 62 245 84.5 123 123 297 35 262 80 113 120 2 626 130 496 170 253 259 609 65 544 172 238 260 2 649 132 517 172 241 253 465 77 388 137 169 175 Gregarina delmasi Tuzet et Rambier, 1953 Solitary individuals Individual TL PL DL PWM DWE DWM 1 471 71 400 156 113 137 2 517 63 454 129 78 148 3 435 125 310 206 109 172 4 485 98 387 129 111 119 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 607 69 538 185 124 188 526 49 477 148 127 183 2 604 82 522 173 79 200 591 92 499 127 114 255 3 578 93 485 181 88 215 589 100 489 133 117 155 Gregarina katherina Watson, 1915 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 104 20 84 28 40 44 111 11 100 30 39 40 2 109 19 90 31 59 60 115 12 103 37 48 49 3 72 12.5 59.5 23 30 32.5 75.5 11 64.5 26.5 33 33 3* 59 8.5 50.5 19 25 25.5 *Data for secondary satellite in association 3 Gregarina barbarara Watson, 1915 Solitary individuals Individual TL PL DL PWM DWE DWM 1 112.5 17.5 95 29.5 66 69 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 108.5 18.5 90 31.5 61 66.5 117 11 106 41 51 53 2 93 19 74 27.5 47 52 97.5 7 90.5 34.5 42 46 3 98 22 76 26.5 50.5 53 98.5 7.5 91 35 40 47 Gregarina steini Berndt, 1902 Solitary individuals Individual TL PL DL PWM DWE DWM 1 103.5 15.5 88 19 26.5 30.5 2 93.5 11 82.5 17 21 26 3 61.5 9 52.5 13 15.5 17 4 128 15.5 112.5 22.5 28 32 5 134 20 114 31 33 39 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 148 23 125 33.5 35 46 135 12 123 33.5 37.5 41 2 141 16.5 124.5 30 35 40 132.5 17.5 115 30 34 40.5 3 147 18 129 31 38.5 43 134 14.5 119.5 32.5 35 39.5 4 150.5 21.5 129 32 39.5 41 108 12 96 26 29 32 5 147 20 127 29 30 37 102 10 92 22.5 27.5 30 6 140 14 126 30 32 44 111 12 99 26.5 28.5 38 Gregarina cuneata Stein, 1848 Solitary individuals Individual TL PL DL PWM DWE DWM 1 248 70 178 38.5 43 47.5 2 158 48.5 109.5 35 42 43 3 57 14 43 16 17 18 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 231 54.5 176.5 42 73 85 229 53 176 73 87 88 2 174 52 122 42 68 75 140 14 126 43 56 58 3 296.5 62.5 234 44 48 60 408 64 346 53 55 55 4 249 57 192 35 37.5 49 233 39 194 31 32 37 5 207 45.5 161.5 37 36 45 270 44 226 35 40 46 Gregarina polymorpha (Hammerschmidt, 1838) Solitary individuals Individual TL PL DL PWM DWE DWM 1 183 32 151 31.5 43 44 2 178 28 150 32 41 44 3 263 40 223 43 55.5 61 4 278 35 243 47 69 73 5 269 24 245 39 61 66 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 256 33 223 41 57 59.5 292.5 25.5 267 51 60 65 2 292 34 258 42 64.5 74.5 309 27 282 55 71 78 3 212 28 184 39 50 53 223.5 25.5 198 43 52 56 4 245 30 215 41 55 60 247 23 224 42 53.5 60 5 351 28 323 43 70 72.5 354 29 325 51 75 77.5 Gregarina curvata (Hammerschmidt, 1838) Solitary individuals Individual TL PL DL PWM DWE DWM 1 456 49 407 53 99 106 2 458 49.5 408.5 58 100 107 Gamocystis tenax Schneider, 1875 Solitary individuals Individual TL PL DL PWM DWE DWM 1* 289 - - - 150 162 2* 284 - - - 136 142 3 287 8 279 36.5 206 206 4 296 8 288 35 214 214 5 325 9 316 40 197 197 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 303.5 8.5 295 40 227 227 345* - - - 178 191 2 261.5 7.5 254 38 164 164 285* - - - 142 151 3 276 7 269 21 143 144 239* - - - 122 122 4 263* - - - 182 187 339* - - - 148 152 5 259* - - - 207 220 345* - - - 169 172 *The cell was not divided into protomerite and deutomerite Gamocystis fimetarii Cordua, 1953 Solitary individuals Individual TL PL DL PWM DWE DWM 1 293 1.5 291.5 14.5 73 73 2 291 1.5 289.5 25.5 90 90 3 324 2 322 19 93.5 95 4 286 2 284 15.5 97 97 5* 275 - - - 81 82 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 315 7 308 37.5 150 154 431* - - - 133 137 2 342 8.5 333.5 33 118 119 384* - - - 105 114 3 318 8 310 38 113 113 425* - - - 106.5 107 4 246* - - - 87 88 235* - - - 57 71 5 295 7 288 35 92 93 309* - - - 78 78 *The cell was not divided into protomerite and deutomerite Hirmocystis polymorpha (Leger, 1892) Assoc 1 Parameter TL PL DL PWM DWE DWM Primite 115 22.5 92.5 33 52 56 Satellite 1 122 11 111 39 55 56 Satellite 2 116.5 14 102.5 34 49 51 Satellite 3 108.5 12 96.5 31 44 44 Satellite 4 118 15 103 32 45 45 Satellite 5 86.5 10.5 76 29 38.5 38.5 Assoc 2 Parameter TL PL DL PWM DWE DWM Primite 297 48 249 88 144 148 Satellite 1 305 34.5 270.5 97 163 163 Satellite 2 342 33 309 109 155 155 Satellite 3 300 29.5 270.5 97 139.5 139.5 Satellite 4 307 35 272 89 136 138 Satellite 5 237 30 207 76 105 105 Satellite 6 178 23.5 154.5 59 76.5 85 Assoc 3 Parameter TL PL DL PWM DWE DWM Primite 102.5 16 86.5 28 48 51 Satellite 1 102 11 91 33 52 52 Satellite 2 117.5 12.5 105 34.5 49 49.5 Satellite 3 104 12 92 31 44 44 Satellite 4 108.5 14.5 94 31 44.5 44.5 Satellite 5 82 11 71 26 35 35 Satellite 6 67 8 59 22 28.5 28.5 Hyalospora hemerobii Geus, 1969 Solitary individuals with epimerite Individual TL EL PL* DL PWM DWE DWM 1 120 29 20 71 33 70 70 2 101 26.5 20 54.5 27 50 50 3 103 33 16 54 26 49 49 4 87.5 31.5 17 39 19.5 36.5 36.5 5 137 31 26 80 35 83 83 Solitary individuals without epimerite Individual TL PL DL PWM DWE DWM 1 104 26 78 37 69.5 71 2 98.5 26.5 72 36 65.5 65.5 3 86 22 64 34.5 59 59 4 266 32 234 60 182 182 5 102.5 25.5 77 37 69 70 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 82 20 62 26.5 47 47 78 14.5 63.5 22 31 32 2 99 24 75 28 60 60 89 13.5 75.5 35 70 70 3 72 19 53 26.5 51 51 83 16 67 25 43 43 Hyalospora psocorum (Siebold, 1B39) Solitary individuals Individual TL PL DL PWM DWE DWM 1 48.5 13 35.5 21 22.5 24 2 49 11.5 37.5 21 16.5 20.5 3 49 12.5 36.5 20 18.5 19.5 4 64.5 12 52.5 18.5 20 23.5 5 69 10.5 58.5 19.5 20.5 24 Euspora fallax Schneider, 1B75 Gamonts in association Assoc Pr Sat TL PL DL PWM DWE DWM TL PL DL PWM DWE DWM 1 492 64 428 92 209 217 454 29 425 102 198 203 2 299 42 257 64 113 128 308 23.5 284.5 71 114 119 3 486 69 417 93.5 236 256 474 39 435 100 260 265 4 513 28 485 100 251 269 222 14.5 207.5 64 80 83.5 5 618 82 536 113 277 293 553 34 519 122 276 276 Leidyana gryllorum (Cuenot, 1B97) Solitary individuals Individual TL PL DL PWM DWE DWM 1 357 53 304 82.5 69.5 95 2 371 48 323 69 55 75 3 441 85 356 96.5 126 129 4 361 62 299 69 60.5 70 5 375 68.5 306.5 68.5 72.5 82 Leidyana oblonga (Dufour, 1B37) Solitary individuals Individual TL PL DL PWM DWE DWM 1 274 40.5 233.5 79.5 120 132 2 287 51 236 71 98 107 Actinocephalus permagnus Wellmer, 1911 Solitary individuals Individual TL PL DL PWM DWE DWM 1 356 48 308 49.5 49 60 2 979 74 905 54 54.5 69 3 754 69 685 56 56.5 78.5 4 470 44.5 425.5 58 62 71.5 5 859 61 798 52 64.5 77 Actinocephalus conicus (Dofour, 1837) Solitary individuals with epimerite Individual TL EL PL* DL PWM DWE DWM 1 314 62 55 197 88 97 112.5 2 274 69 54 151 67.5 75.5 79 3 448 98 69 281 148 171 194 4 259 51 50 158 73 75.5 84 5 266 53.5 55.5 157 68 73 81 Solitary individuals without e pimerite Individual TL PL DL PWM DWE DWM 1 679 154 525 232 204 257.5 2 317 77.5 239.5 129 150 159 3 791 181.5 609.5 239.5 222 285 4 762 127 635 165 234 291 5 321 86 235 130 142 174