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ISSN 1330-0520 UDK 595.76.581.96(497.5)

VOL. 9 No 2


ZAGREB June 30, 2000

original scientific paper / izvorni znanstveni rad


Dipartimento di Zoologia e Antropologia Biologica dell’Università, Via Muroni 25, 07100 Sassari, Italy

Museo Regionale di Scienze Naturali, Via Giolitti 36, 10123 Torino, Italy


Department of Zoology, Croatian National History Museum, Demetrova 1, HR–10000 Zagreb, Croatia

Casale, A., Giachino, P. M. & Jal`i}, B.: Croatodirus (nov. gen.) bozicevici n. sp., an enigmatic new leptodirine beetle from Croatia (Coleoptera, Cholevidae). Nat. Croat., Vol. 9, No. 2., 83–92, 2000, Zagreb. Croatodirus, new genus, bozicevici new species (Coleoptera, Cholevidae) are described from the cave Cavern in U~ka Tunnel, Mt. U~ka (Monte Maggiore) near the town of Rijeka (Croatia). The new taxon, provisionally attributed to the phyletic lineage of Antroherpon, is however markedly isolated owing to the following peculiar combination of morphological features: 1) antennal insertion on the posterior 4th of the head; 2) antennomere 1 slightly longer than antennomere 2; 3) pterothorax not pedunculate, mesosternum not carinate; 4) mesocoxal cavities contiguous; 5) tarsal claws widened; 6) protibiae without external apical comb of spines and without external spurs; 7) meso- and metatibiae furnished with inner, unified spurs; 8) aedeagus medium sized, with inner sac without sclerotized pieces; and 9) female stylomeres and spermatheca fully atrophied. The systematic position and the peculiar ecology of this new, exceptional troglobiontic beetle are illustrated and discussed. Key words: Croatodirus, bozicevici, new genus, new species, Coleoptera, Cholevidae, systematics Casale, A., Giachino, P. M. & Jal`i}, B.: Croatodirus (nov. gen.) bozicevici n. sp., novi neobi~ni leptodirski tvrdokrilac iz Hrvatske (Coleoptera, Cholevidae). Nat. Croat., Vol. 9, No. 2., 83–92, 2000, Zagreb. Novi rod Croatodirus i nova vrsta bozicevici (Coleoptera, Cholevidae) opisani su iz {pilje Kaverna u tunelu U~ka u U~ki (Monte Maggiore) pokraj Rijeke (Hrvatska). Nova svojta, provizorno pripojena fileti~koj liniji Antroherpon, je morfolo{ki ipak zna~ajno izolirana zbog neobi~ne kombinacije sljede}ih morfolo{kih osobina: 1) ticala su smje{tena na stra`njoj ~etvrtini glave; 2) prvi ~lanak ticala je neznatno du`i od drugog ~lanka; 3) pterotoraks nije ispup~en, mezosternum nije grebenast; 4) mezokoksalne {upljine se dodiruju; 5) tarzalne pand`ice pro{irene; 6) protibije bez vanjskog
Croatian Natural History Museum, Demetrova 1, Zagreb, Croatia

Penultimate labial palpomere long. Klju~ne rije~i: Croatodirus. pubescent. will be object of a subsequent contribution. just discovered in nearby areas. = section Antroherpona of JEANNEL. basal margin as wide as the base of elytra. Type species: Croatodirus bozicevici n. Furthermore. and longer than antennomere 2. It was discovered during speleological explorations performed by the Department of Zoology. subrectangular in shape. and NEWTON. for which a new.. 7) mezotibije i metatibije imaju jednake unutarnje trnove. and is well known today for its high specific diversity and the exceptional specialization of many of the taxa. s unutra{njom vre}icom bez sklerotiziranih dijelova. very long. GEN. however. A. Croatian Natural History Museum in Zagreb. nova vrsta. with its maximum width just before the middle. Cholevidae. bozicevici. medium long pubescence. and demonstrate that our knowledge of the subterranean fauna of the Dinaric region is still far from comlplete. ultraspecialized leptodirine beetle.: Croatodirus (nov. Some other.. without occipital carina. antennomere 11 shorter than antennomere 10. Pronotum large. vr{nog ~e{lji}a igala i bez vanjskih trnova. et al. ovate elytra. apical palpomere very short. they will allow a re-examination of some morphological characters currently emphasized in reconstructed phylogenies of this group. sp. 1976. 1991 and GIACHINO et al.. free. 9) stilomere `enki i spermateka potpuno atrofirale. in older entomological literature). Head elongate. 1924. with hind angles obtuse but evident. Antroherponina of GUÉORGUIEV. large. sistematika INTRODUCTION The subterranean fauna of Croatia has been the object of extensive investigations in the last two centuries (NONVEILLER.) bozicevici n. slightly sinuated basally. A genus of medium sized. subrectangular pronotum. 8) edeagus srednje veli~ine. sp. very interesting troglobitic Leptodirinae. 1998. izuzetnog troglobiontskog kornja{a i o njegovoj neobi~noj ekologiji. U radu se daje opis i raspravlja o sistematskom polo`aju ovog novog.84 Casale. Coleoptera. isolated genus is proposed. eyeless. gen. Several troglobitic organisms. and will furnish further data on of the hypogean cholevids of the region. 1998) with a markedly pholeuonoid body. infraflagellate Leptodirini (sensu CASALE et al. Antennae inserted on the posterior 4th of head. CROATODIRUS NOV. lateral sides regularly rounded anteriorly. antennomere 1 slightly widened at apex.. and exceeding the elytron apex. have been discovered and described only in recent years. 1999). The present contribution deals with a new. clypeus and labrum with dense. and antennae longer than the body length. similar in both sexes. and comes from a carefully explored coastal area close to the town of Rijeka (known as »Fiume«. . . novi rod.

2000 85 Fig.Nat. . holotypus m: habitus. Scale: 1 mm. gen. Croat. 9(2). Vol. Croatodirus (nov.) bozicevici n. 1.. sp.

with three basal tarsomeres dilated. similar in both sexes. Scale: 0. Legs very long and slender.. Aedeagus (Figs. 6) female genital segment. A.: Croatodirus (nov. 2–6. parts of holotypus and one paratypus: 2) aedeagus lateral view. 4) left paramere. each furnished with three short apical setae. Microsculpture not forming transversal rows. Elytra elongate ovate.1 mm. ventrite only slightly sclerotized and furnished with a group of short. et al.) bozicevici n. . Male genital segment reduced in size. Female genital segment (Fig. Male protarsi 5-segmented. Mesosternal carina absent. thin pubescence. Inner sac unarmed. 2–5) medium sized. thickened setae on each side.. sp. disc with short. gen. mesocoxal cavities contiguous. regularly arcuate. basal lamina of median lobe very arcuate. with femora thickened basally. Figs. Ovipositor and spermatheca fully atrophied. . lateral view. parameres as long as the median lobe. dorsal view. attenuate at apex. tarsal claws long and slyghtly widened. stout. Protibiae widened at apex. sutural stria absent. without apical comb and outer and inner spurs. sp. 6) membranous. with a little evident ventral carina. gen.86 Casale. Pterothorax not pedunculate. Croatodirus (nov. with inner unified apical spur and apical row of apical spines. 3) aedeagus dorsal view. curly shaped.) bozicevici n.. 5) right paramere.

75 m. 15.52. Torino). Aedeagus (Figs. Mt.02. respectively). disc subconvex. in dorsal view. Mouth parts adapted to a »Hadean« way of life. 12. 6. Croatia. cave Cavern in U~ka Tunnel Type material: Holotypus male. 1999. and remnants of 1 specimen. 6 females.80 m. 1999. 13.32.. 5. Giachino. 2–5) medium sized. short. Median lobe. 13. Antennomere ratio: HT m: 4.40 PT f: 4. 3. Croatia. decumbent pubescence. slightly shorter in the female (ratio ML / MW: 1. Coll. 6. (Coll. 10. Paratypi: 2 males.14. U~ka.35. 9(2).67 f).69. Rijeka.48–3. Rijeka. Jal`i} leg. B.35. cave Cavern in U~ka Tunnel. ultraspecialized leptodirine beetles. similar in size and length. U~ka. 1.71. 1924.52 mm. acuminate apically. 27. Antennae very long and slender (ratio prothorax + elytra / antennae: 0. short pubescence. NONVEILLER & PAVI^EVI]. to filtering water and organic matter (JEANNEL. 11. who did a lot of resarch on this cave and many others in the Dinaric karst. Mt. sp. 9. Sep.56. in lateral view arcuate. 1 female. 4. 1. 8. with maximum width at middle.09 m.e. not retractile. covered by dense. 12. subtruncate at apex.15 Pronotum large in size.77.70 ff). integument opaque. 12. Head elongate. covered by dense. Jal`i} leg. 11. subrectangular. Rijeka. Rijeka. U~ka. 6. Croatia.78. longer than wide (ratio ML / MW: 1. Croat. 2000 87 DERIVATIO NOMINIS Croatodirus: epithet composed of the names »Croatia« and »Leptodirus«. 0. 6. Casale. Legs: see description of the genus. Elytra elongate ovate. Mt. Parameres each furnished with three short setae (one apical. . the others pre-apical. 1996. 5. 14.17. dorsally depressed in the apical third. Specific epithet The new species is dedicated to the retired geologist and speleologist Sre}ko Bo`i~evi} PhD.79. Torino. Colour dark reddish.34. 15. DESCRIPTION A medium sized (TL: mm 3. cave Cavern in U~ka Tunnel. 14. Disc very convex.59. (Croatian Natural History Museum).95. Croatodirus bozicevici n. Typ. Loc. Jal`i} leg. highly specialized leptodirine beetle. B. 1999). Mt. Oct. anophthalmous. Apex fully covering the pygidium. slightly constricted towards the base.16 f). stout. cave Cavern in U~ka Tunnel.68–3.32. the latter being a well known genus of troglobitic. CASALE & JAL@I].74. Oct. B.97. pholeuonoid. i.Nat.80 f).37. Vol. without occipital carina. 1988. pubescent.: Croatia. U~ka. 4.

: Croatodirus (nov. measured in September 26th. The water temperature. an entrance permit must be obtained. 7) in 1979.88 Casale. Groundwater tracing proved that the waters from the cavern supply the coastal spring from the settlement of I~i}i to the settlement of Medveja. The flow velocities measured in the cavern range from 10 to 30 l/s. A natural cave opening was not discovered. Map of Croatia with the position of cave Cavern in U~ka Tunnel (•). sp. Since it is located within a water supply extraction site and along the route of the tunnel. . The groundwater flow is formed by surface waters drainomg from the region of Crkveni Vrh peak. turbulent flow also occurs. DISTRIBUTION. The reverse faults and the overthrusting of Cretaceous carbonate rocks into impermeable flysch deposits were a prerequisite for the formation of the cavern. Fig.. caused the formation of large caves. . Periodically. The origin of the cave is a consequence of tectonic events within the U~ka massif. along the areas of contact of the flysch rocks and the limestone. so that access to the cavern is possible only through an artificial shaft. 7. Speleological investigations allowed the discovery of 1490 m of galleries with an elevation difference of 135 m (Fig. et al. 8).5 °C.) bozicevici n. gen. ECOLOGY The cave Cavern U~ka Tunnel was discovered after rock blasting during the excavation of the U~ka Road Tunnel near Rijeka (Fig. 1996. the cavern is geologically young and still under the influence of intensive geological transformation. The movement of water. In a genetic sense.. was 8. A.

which does not contain any visible traces of human activity. The faunistic findings are located in the area after the siphon. Vol. An investigation into the subterranean fauna was carried out twice. A stream flows through the chamber over rocks that are located on its floor. and October 15th. At this location. during geological speleological studies on September 26th. 2000 89 Fig. . Part of the plan of cave Cavern in U~ka Tunnel. Parts of the channel are covered by dust and soot from blasting. and therefore can be considered as intact. Only in the upper part of the chamber do sandy and muddy sediments occur along the banks of a somewhat large lake/pool. 1996.Nat. Sampling was made in the chamber after the siphon named the Nova dvorana (New chamber). Croat. In the area that leads to the siphon no animal organisms were found: this is probably a temporary situation caused by the tunnel-building work. 8. 9(2). 1999. there is intensive seepage of water from the chamber ceiling.

1959 and Niphargus krameri Schellemberg. and suggest a particular. Furthermore. 1935.. . gen. Deelemania pretneri Perreau. 1998) as a whole. 6) tarsal claws dilated (less dilated however than in Antroherpon. et al. the species Typhlotrechus bilimeki istrus (G. sp. is a character already reported for other subterranean cholevid beetles (GIACHINO et al. 1998. the following crustacean species were collected and identified: the terrestrial species Titanethes dahli Verhoeff. sp. to Radziella styx Casale & Jal`i}. on a global scale. however. adapted to life in deep hypogean compartments: in particular. 1988. absent in Antroherpon). 1998). superficially similar to other leptodirine taxa of Dinaric Alps. A more careful examination shows. many peculiar morphological characters. Finally. and the stygobian species Monolistra bericum hadzii Sket. 26). from Durmitor (Crna Gora).) bozicevici n. In particular: 1) pterothorax not pedunculate. 3) mesocoxal cavities contiguous.. Carabidae) was also found. Beside these. 1926) (Coleoptera. 5) protibiae without apical comb of spines and without external spurs. It should belong to the phyletic lineage of Antroherpon (in the sense of the authors) owing to the following features: 1) antennae inserted in the posterior 4th of the head. and carefully weighing morphological features that are markedly convergent in different subterranean taxa adapted to the same way of life. Müller. from Biokovo (Croatia).: Croatodirus (nov.90 Casale. Reduced spermatheca. the full atrophy of female stylomeres and spermatheca is peculiar to Croatodirus. from Sana Valley (Bosnia). among others – could be polyphyletic assemblages of taxa. A.. Such an examination of taxonomic groups. now seems necessary for some phyletic lineages which – as hypothised about some trechine Carabids by SCIAKY & VIGNA TAGLIANTI (1990) and CASALE & JAL@I] (1999). RELATIONSHIPS Its general features and degree of specialization make Croatodirus (n. however. All the characteristics illustrated and discussed above demonstrate the need for a new and original phylogenetic analysis of the Leptodirinae (sensu NEWTON. 4) femora thickened basally.) bozicevici n. which isolate the new »infraflagellate« taxon from the monobasic genera cited above (»Theleomorphes« sensu GIACHINO et al. Leptodirus-like. going beyond traditional classifications. and from all other dinaric leptodirine genera known so far. 1926. 1999. and Tartariella durmitorensis Nonveiller & Pavi~evi}. unknown way of fecundation and oviposition. and in other troglobitic cholevids specialized to life on stalagmitic walls). linked . narrowed apically. Radziella. Fig. Some other characteristics. on the other hand. and meso-and metatibiae each provided with inner unified spur and apical row of spiniform setae (the latter. 3) aedeagus medium sized. 1999. 2) antennomere 1 longer than antennomere 2. Most specimens of the new beetle species were found on the stalagmitic encrustations along the stream.. suggest Croatodirus also has relationships with the phyletic lineage of Leptodirus (of the authors). gen. 2) male genitalian segment very reduced in size. such as in Leptodirus (less reduced in the examined Antroherpon species).

Jugoslavia. NEWTON.. P. GIACHINO. D. M. JEANNEL. A. P. B. 1999: The Pioneers of the research on the Insects of Dalmatia. Arch.Nat. Carabidae. G.. 2000 REFERENCES CASALE. Bull.. Croatian Natural History Museum. reg. in which further new. sp. Nat. A. F. exp. ital. & JAL@I]. 63. current classification and generic catalog of world Leiodidae (including Cholevidae). ACKNOWLEDGEMENTS We are indebted to the geologists Sre}ko Bo`i~evi} and Mladen Kuhta (Institut of Geology. Florence. & CASALE.. Mus. Catopidae) from the Dinaric Region. entomol. 1988: Radziella (new genus) styx n. who enabled us to join their team during the speleological investigations of the cave Cavern in U~ka Tunnel. Atti XVI Congr. M.. P.. Bari-Martina Franca (TA).. In: GIACHINO. & JAL@I]. V. Croat. D. GIACHINO. 1976: Recherches sur la taxonomie. We also express our thanks to Sanja Gottstein. 349–358. Boll. 1996. 1998: Phylogenetic problems. Atti Mus. PERREAU.. A. sp. 1991: Brevi considerazioni per una sistematica filogenetica dei Bathysciinae (Coleoptera: Cholevidae). Torino. B. XX International Congress of Entomology. S..). Leiodidae). Phylogeny and Evolution of Subterranean and Endogean Cholevidae (=Leiodidae Cholevinae). Sci. S. Such an analysis will be presented in a subsequent contribution. ultraspecialized troglobitic cholevids from Croatia. 857–865. Sci. a new species of eyeless Trechinae beetle from Gorski Kotar (Coleoptera. Torino. (Paris). a new exceptional troglobitic Bathysciinae (Coleoptera. 1998: Major questions in the phylogeny and biogeography of Cholevidae (Coleoptera). Proceedings of a Symposium (30 August. B. 1–436. M. XX International Congress of Entomology.. naz. nat. CASALE. VAILATI. 9(2). & VAILATI. génér. Sci. A.. 104(4). (eds. GUÉORGUIEV. NONVEILLER. ent. 2000 91 more by similar adaptive features than by real phyletic relationships. M.. will be described. Dissert. CASALE. 137–145. 179–209. as a result of independent. Soc. Acad. Artium Slov.. & PECK. 1999: Nouveaux genres et nouvelles espèces de Leptodirini (Coleoptera. Fr. Florence. A. unrelated epigean ancestors.. Vol. Italy). In: GIACHINO. 6(2). Zagreb). reg. Phylogeny and Evolution of Subterranean and Endogean Cholevidae (=Leiodidae Cholevinae). la classification et la phylogénie des Bathysciinae. Zagreb. & PECK.. R.. Proceedings of a Symposium (30 August. Sci. M. 8(2).. Razp. B. 390 pp. 41–178. 1996. . who identified the Crustacea collected. 399–406. Trechini). Croat. nat. Received March 1. (eds. B. P. Italy). 1924: Monographie des Bathysciinae. Atti Mus..). reg. zool. Torino. 1999: Croatotrechus (new genus) tvrtkovici n. 91–147 (1–59). just discovered. heterochronic phases of colonization of the subterranean environment by different.. Biospeologica L. with emphasis on the subfamily Leptodirinae. nat. 19(4)..

17.. NONVEILLER. A.. A. et n. Mem. 1982. sp.92 Casale. D. Fr. Trechinae).. 1990: The genus Lessinodytes Vigna Taglianti. G. Biospéol.. sp. ent.. Soc. 315–326. . troisième Leptodirini de la chaîne Dinarique à moeurs hadésiennes (Coleoptera. a biogeographical and systematic puzzle (Coleoptera.. gen. 104(4). SCIAKY. & VIGNA TAGLIANTI. Carabidae. gen. & PAVI^EVI]. 1999: Tartariella durmitorensis n. Bull.: Croatodirus (nov. 169–173.) bozicevici n. et al. R. Leiodidae). .