Two new endemic species of the genus Omphreus (Carabidae: Coleoptera) from Montenegro

Biologia 63/3: 402—406, 2008 Section Zoology DOI: 10.2478/s11756-008-0063-6

Two new endemic species of the genus Omphreus (Carabidae: Coleoptera) from Montenegro ´ ´1, Wolfgang Waitzbauer2, Pamela Zolda2, Miloje Brajkovic ´ 1, Sre´cko Curči c ´ ´1, Nastas Ilic ´3 & Božidar P.M. Curči ´1 Rajko Dimitrijevic c 1

Institute of Zoology, Faculty of Biology, University of Belgrade, Studentski Trg 16, 11000 Belgrade, Serbia and Center for Biospeleology of Southeast Europe, Resavska 53, 11000 Belgrade, Serbia; e-mails: [email protected], [email protected] 2 Department of Conservation Biology, Vegetation- and Landscape Ecology, Faculty of Life Sciences, University of Vienna, Althanstrasse 14, A-1090 Vienna, Austria; e-mails: [email protected], [email protected] 3 Faculty of Sport and Physical Education, University of Belgrade, Blagoja Parovi´ca 156, 11000 Belgrade, Serbia; e-mail: [email protected]

Abstract: Two new species, viz. Omphreus (Omphreus) prekornicensis sp. n. (from Medjedje Peak, Mt. Prekornica, C Montenegro) and Omphreus (Omphreus) bjelasicensis sp. n. (from Biogradska Gora National Park, Mt. Bjelasica, E Montenegro), are described and diagnosed. Male and female genitalia and other taxonomically important characteristics are illustrated. The new species are clearly distinct from their closest congeners and represent endemic relicts inhabiting limited regions of the Dinaric mountain system in Montenegro. Key words: Carabidae; Omphreus; new species; soil fauna; Montenegro

Introduction

Material and methods

The genus Omphreus Dejean, 1828 comprises three subgenera (Neomphreus Winkler, 1933, Omphreus s. str., and Paromphreus Ganglbauer, 1887), 14 species, and 13 subspecies, which are distributed both on the Balkan Peninsula and in Asia Minor (Trautner & Geigenm¨ uller 1987; Bousquet 2003). All Omphreus taxa are endemic and most are montane to alpine forms living in forests (under rocks and bark), but also under stones beyond the timberline. These carabids are large in size (16–28 mm), black, mostly very rare, and recognizable both by their habitus and by the long shaft-like first antennomere. The territory of Montenegro is inhabited by the following Omphreus taxa: Omphreus (Neomphreus) apfelbecki meridionalis Winkler, 1933 (from Kremeni Do near the village of Gornje Stravče, Mt. Žijovo, SE Montenegro, and the village of Lijeva Rijeka, Mt. Žijovo, SE Montenegro), Omphreus (Omphreus) morio beckianus Ganglbauer, 1888 (from the region of the border with Bosnia-Herzegovina), Omphreus (O.) morio malissorum Winkler, 1933 (from Mt. Prokletije, E Montenegro), Omphreus (O.) morio morio Dejean, 1828 [from the regions of Njeguši (Mt. Lov´cen) and Krivošije (Mt. Radostak), SW and S Montenegro], and Omphreus (O.) wohlberedti Winkler, 1933 (from Virpazar near the Skadar Lake, S Montenegro) (Winkler 1933; Drovenik & Peks 1999; Bousquet 2003).

Two field trips organized by one of the coauthors of the present study (Prof. Dr. Nastas Ili´c) in C and E Montenegro during 2001 and 2002 resulted in the discovery of two new species of the genus Omphreus: Omphreus (Omphreus) prekornicensis sp. n. and Omphreus (Omphreus) bjelasicensis sp. n. The specimens of the new species were collected by hand and with Barber vinegar traps. The type specimens were analyzed in laboratories of the Department of Conservation Biology, Vegetation- and Landscape Ecology, Faculty of Life Sciences, University of Vienna; there they were dissected, thoroughly studied, and illustrated. Dry specimens were stuck on rectangular paper labels, while the male and female genital structures were stuck on separate triangular paper labels and deposited with the specimen from whom they were extracted. All important morphological characteristics were studied with Wild M3C and Nikon SMZ-U binocular stereomicroscopes. A Nikon Coolpix 4500 digital camera with special monitor attached has been used for taking the photographs.

c 2008 Institute of Zoology, Slovak Academy of Sciences


´ Omphreus (Omphreus) prekornicensis S. Curči´ c sp. n. (Figs 1–5) Description. Large-sized. Body length: 19.15–22.54 mm (average 20.77 mm). Body elongated; elytra ovoid (Fig. 1). Body color black, mouthparts, apical antennomeres, and tarsi black-brownish. Tegument shiny, except for slightly rough elytra.

New species of the genus Omphreus

Figs 1–5. Omphreus (O.) prekornicensis sp. n.: 1 – male (holotype), habitus (dorsal view); 2 – male (holotype), aedeagus (dorsal view); 3 – male (holotype), aedeagus (lateral view); 4 –female (paratype), genitalia; 5 – male (holotype), abdominal sternite IX (urite). Scales 1 mm.

Head rounded, somewhat longer than wide, its width/length ratio: 0.89 (0.81–1.00), shorter (head/ pronotum length ratio: 0.66) and narrower (head/pronotum width ratio: 0.68) than pronotum (Fig. 1). Head below eye level visibly constricted. Labrum wide, medially with rounded convexity, bearing four setae. Epistome large, with concave fore border, bisetose. Vertex and occiput relatively wrinkled, without fine punctuation. Frontal foveae deep and relatively long. Gula with two setae. Mandibles long and sickle-shaped, widened proximally. Labial palpomeres with first article shorter than second, without setae, second and third articles widened distally, ultimate article more densely setose. Maxillary palpomeres with two distalmost articles densely pubescent. Average antennal length: 12.11 mm (10.23–13.00 mm). Antennae pubescent from fourth antennomere. Ratio of length antennae/body: 0.57 (0.53– 0.62). First antennomere sharply widened distally, with sparsely long setae apically, shorter than the following three antennomeres together. Antennomere II shorter than antennomere III. Pronotum sub-bell-shaped, relatively elongated, its width/length ratio: 0.87 (0.81–0.93), slightly convex

403 apically and basally (lateral view), fore angles prominent and rounded apically, hind angles somewhat obtuse with rounded top (Fig. 1). Lateral margins clearly convex anteriorly and concave posteriorly. Pronotum widest slightly before end of its fore third. Anterior and especially posterior margins concave. Lateral margins well-expressed, wide, with four anterior setae, one median seta, and one posterior seta. Lateral furrows relatively wide and deep. Median furrow almost as long as length of pronotum. Basal foveae deep and long, somewhat shorter than half of pronotal length. Elytra oval, elongated, slightly convex (lateral view), with arcuated lateral sides, their width/length ratio: 0.57 (0.55–0.60) (Fig. 1). Elytral width: 6.95 mm in males, 7.69 mm in females. Elytral striae weakly expressed, but visible. Scutellar striae developed, scutellar punctures present. Elytra widest somewhat after level of their half-way point. Seventh interstria with one or two basal setiferous punctures (situated close to sixth stria), four or five median setiferous punctures (situated in seventh interstria), and two apical setiferous punctures (situated in seventh stria). Setiferous punctures very large and deep. Striae fine, very slightly punctuated. Shoulders weakly obtuse. Umbilicate series regular, its setae densely distributed. Intervals of elytra flattened. Two fore protarsomeres in males dilated; metacoxae long and rounded. Tarsal claws long, glabrous, without teeth (Fig. 1). Aedeagus long, regularly curved laterally, widened sub-apically (in both dorsal and lateral view), with rounded triangular top. Paramerae long and moderately wide. Basal bulb not widened distally (Figs 2, 3). Female genitalia (Fig. 4): gonocoxites IX welldeveloped, more curved, thick, distally moderately narrowing, apically rounded, basally joined with massive gonosubcoxites IX. Male abdominal sternite IX (urite) well-developed, sub-triangular (Fig. 5). Material examined. Holotype (male): Montenegro, Mt. Prekornica, village of Jugovi´ci, Medjedje Peak, 900 m a.s.l. near Nikši´c (19◦ 3 E, 42◦ 40 N), 28.VIII.–9.IX.2002, leg. Z. , 1
, same locality as for holotype, Zlati´c. Paratypes: 8 10.VII.–28.VII.2001, leg. N. Ili´c. The type series is deposited in the collection of the Center for Biospeleology of Southeast Europe, Belgrade, Serbia (CBSEE-06/96-105).

Etymology. The new species is named after Mt. Prekornica in C Montenegro, its terra typica. Differential diagnosis. The newly described species is compared below with the phenetically close species Omphreus (O.) morio, O. (O.) wohlberedti, and O. (O.) bjelasicensis sp. n. The new species is considerably longer than O. (O.) morio (20.77 mm vs. 15.00–20.00 mm). Elytral striae are more impressed in O. (O.) morio than in O. (O.) prekornicensis sp. n. Elytra are narrower in O. (O.) morio than in the new species considered. The

404 body plan and degreee of specialization point to certain similarities between Omphreus (O.) prekornicensis sp. n. and some subspecies of O. (O.) morio from Montenegro, viz., O. (O.) morio beckianus, O. (O.) morio morio, and O. (O.) morio malissorum. A more careful examination, however, reveals several clear differences of character states between these four taxa. Thus, Omphreus (O.) prekornicensis sp. n. differs from O. (O.) morio beckianus in body length (19.15–22.54 mm vs. 16.00–18.00 mm), the antennae/body length ratio (0.57 vs. 0.55), presence/absence of the median labrum convexity (present vs. absent), the head width/length ratio (0.89 vs. 1.03), the head/pronotum length ratio (0.66 vs. 0.61), the pronotum width/length ratio (0.87 vs. 0.92), shape of both fore and hind pronotal angles (prominent vs. not prominent), form of the lateral pronotal margins (anteriorly clearly convex, posteriorly strongly concave vs. anteriorly slightly convex, posteriorly weakly concave), shape of the elytra and their lateral margins (oval, with well rounded lateral margins vs. narrow, with slightly rounded lateral margins), the elytral width/length ratio (elytra more than two thirds longer than broad vs. elytra about two thirds longer than broad), the elytra width/length ratio (0.57 vs. 0.60), and the level of maximum elytral width (somewhat after mid-elytra length vs. at the level of midelytra length). Omphreus (O.) prekornicensis sp. n. differs from O. (O.) morio morio in body length (20.77 mm vs. 20.00 mm), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), form of the elytra (wider, ovoidly oval vs. thinner, oval), and the elytral width/length ratio (elytra more than two thirds longer than broad vs. elytra almost two times longer than broad). Finally, O. (O.) prekornicensis sp. n. differs from O. (O.) morio malissorum in body length (19.15–22.54 mm vs. 17.00– 18.00 mm), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), shape of the pronotum (strongly widened vs. less widened), form of the elytra (wider, ovoidly oval vs. narrower, oval), and the elytral width/length ratio (elytra more than two thirds longer than broad vs. elytra almost twice as long as broad). Omphreus (O.) prekornicenis sp. n. clearly differs from Omphreus (O.) wohlberedti in body size (20.77 mm vs. 26.00 mm), size of the head (proportionally larger vs. proportionally smaller), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), the pronotum width/length ratio (pronotum much longer than wide vs. pronotum somewhat longer than wide), presence/absence of a row of points in the second interstrial spaces (absent vs. present), shape of the elytra (ovoidly oval, with well rounded lateral margins vs. oval, with weakly rounded lateral margins), the elytral width/length ratio (elytra more than two thirds longer than broad vs. elytra more than twice as long as broad), and development of brush-

´ ´ et al. S. Curči c like setation on the ventral side of the two fore male protarsomeres (well-developed vs. developed to a lesser extent). Omphreus (O.) prekornicensis sp. n. clearly differs from Omphreus (O.) bjelasicensis sp. n. in average body size (20.77 mm vs. 18.67 mm), the head width/length ratio (0.89 vs. 0.935), antennal length (12.11 mm vs. 10.62 mm), length of the first antennomere (much shorter than antennomeres II–IV together vs. slightly shorter than antennomeres II–IV together), the antennomere II/III length ratio (0.94 vs. 1.00), the head/pronotum length ratio (0.66 vs. 0.64), the degree of head constriction below eye level (more constricted vs. less constricted), surface of the vertex and occiput (relatively wrinkled vs. less wrinkled), position of a weak pronotal convexity in lateral view (apically and basally vs. anteriorly), form of the fore pronotal angles (more prominent vs. less prominent), shape of the lateral pronotal margins (very convex anteriorly, slightly concave posteriorly, sub-parallel basally vs. less convex anteriorly, more concave posteriorly, divergent basally), width of the lateral pronotal margins (wider vs. narrower), appearance of the lateral pronotal furrows (deep and wide vs. somewhat shallower and narrower), maximum width of the pronotum (somewhat before the end of its fore third vs. at the end of its fore third), the elytra width/length ratio (0.57 vs. 0.59), elytral width (6.95 mm in males, 7.69 mm in females vs. 6.41 mm in males, 6.42 mm in females), maximum width of the elytra (somewhat behind their half-way point vs. at the level of their half-way point), shape of the elytra (ovoidly oval vs. regularly oval), impression of the elytral striae (more expressed vs. less expressed), presence/absence of scutellar punctures (present vs. absent), position of median elytral setiferous punctures in seventh interstrial spaces (usually not close to seventh striae vs. close to seventh striae), shape of the apex of the median lobe (rounded vs. pointed), width of the paramerae (narrower vs. wider), appearance of the aedeagus in lateral view (with a wider and more curved median lobe and less massive basal bulb vs. with a narrower and less curved median lobe and more massive basal bulb), shape of gonocoxites IX (more curved vs. less curved), form of the male abdominal sternite IX (urite) (larger, with thicker margins vs. smaller, with narrower margins), and length of the aedeagus (longer vs. shorter) (present study) (Figs 1–10). Bionomy and distribution. The new species inhabits high-altitude sites on Mt. Prekornica, especially epigean and endogean habitats on Medjedje Peak. These soildwelling beetles are predators, their diet restricted mostly to other insects and earthworms. The species represents an endemic relict form, probably of Tertiary origin. ´ Omphreus (Omphreus) bjelasicensis S. Curči´ c et Ili´ c sp. n. (Figs 6–10) Description. Large-sized. Body length: 17.31–19.77

New species of the genus Omphreus

Figs 6–10. Omphreus (O.) bjelasicensis sp. n.: 6 – male (holotype), habitus (dorsal view); 7 – male (holotype), aedeagus (dorsal view); 8 – male (holotype), aedeagus (lateral view); 9 –female (paratype), genitalia; 10 – male (holotype), abdominal sternite IX (urite). Scales 1 mm.

mm (average 18.67 mm). Body elongated; elytra oval (Fig. 6). Body color black, mouthparts, apical antennomeres, and tarsi black-brownish. Tegument shiny, except slightly rough elytra. Head rounded, somewhat elongated, its width/ length ratio: 0.935 (0.90–0.975), shorter (head/pronotum length ratio: 0.64) and narrower (head/pronotum width ratio: 0.69) than pronotum (Fig. 6). Head below eye level less constricted. Labrum wide, medially with a rounded convexity, bearing four setae. Epistome large, with concave fore border, bisetose. Vertex and occiput moderately wrinkled, without fine punctuation. Frontal foveae deep and relatively long. Gula with two setae. Mandibles long and sickle-shaped, widened proximally. First article of labial palps shorter than second, without setae. Second and third articles of labial palps widened distally, the ultimate one more densely setose. Two distalmost articles of maxillary palps densely pubescent. Average antennal length: 10.62 mm (9.69–12.00 mm). Antennae pubescent from fourth antennomere. The antennae/body length ratio: 0.57 (0.52–0.63). First antennomere sharply widened distally, with sparsely distributed long setae apically, slightly shorter than the following three antennomeres together. Antennomere II as long as antennomere III.

405 Pronotum sub-bell-shaped, long, its width/length ratio: 0.87 (0.84–0.90), slightly convex anteriorly (lateral view); its fore angles not too prominent, rounded apically; hind angles somewhat obtuse, with rounded top (Fig. 6). Lateral margins arcuated anteriorly, concave and slightly divergent posteriorly. Pronotum widest in end of its fore third. Anterior and especially posterior margins concave. Lateral margins wellexpressed, wide, with four anterior setae, one median seta, and one posterior seta. Lateral furrows relatively wide and deep. Median furrow almost as long as pronotum itself. Basal foveae deep and long, somewhat shorter than half of pronotal length. Elytra oval, relatively long, slightly convex (lateral view), with arcuated lateral sides, their width/length ratio: 0.59 (0.57–0.62) (Fig. 6). Elytral width: 6.41 mm in males, 6.42 mm in females. Elytral striae weakly expressed. Scutellar striae present, but paired scutellar punctures absent. Elytra widest in middle. One setiferous puncture situated in seventh interstrial spaces basally (close to sixth striae), four or five median setiferous punctures situated in seventh interstrial spaces (close to seventh striae), and two setiferous punctures situated on seventh striae apically. Setiferous punctures very large and deep. Striae fine, shallower than in O. (O.) prekornicensis sp. n., very slightly punctuated. Shoulders weakly obtuse. Umbilicate series regular, its setae densely distributed. Elytral intervals flattened. Two fore protarsomeres in males dilated; metacoxae long and rounded. Tarsal claws long, glabrous, without teeth (Fig. 6). Aedeagus long, regularly curved laterally, widening sub-apically (in both dorsal and lateral view), with dragged pointed triangular top. Parameres long and wide. Basal bulb not widened distally (Figs 7, 8). Female genitalia (Fig. 9): gonocoxites IX relatively thick and long, less curved, somewhat narrowing distally, each with rounded apex, basally joined with massive gonosubcoxites IX. Male abdominal sternite IX (urite) well-developed, sub-triangular (Fig. 10). Material examined. Holotype (male): E Montenegro, Mt. Bjelasica (Biogradska Gora National Park) near Mojkovac (19◦ 38 E, 42◦ 54 N), VIII.2002, leg. N. Ili´c. Paratypes: 6 , 2

, same data as for holotype. The type series is deposited in the collection of the Center for Biospeleology of Southeast Europe, Belgrade, Serbia (CBSEE-06/106-114).

Etymology. The new species is named after Mt. Bjelasica in E Montenegro, its terra typica. Differential diagnosis. The newly described species is compared below with the phenetically close species Omphreus (O.) morio, O. (O.) wohlberedti, and O. (O.) prekornicensis sp. n. Omphreus (O.) bjelasicensis sp. n. and O. (O.) morio are different in shape of the posterior part of the lateral pronotal margins (divergent posteriorly vs. somewhat narrowed or almost sub-parallel, never divergent posteriorly). Also, elytral striae are more im-

´ ´ et al. S. Curči c

406 pressed in O. (O.) morio than in O. (O.) bjelasicensis sp. n. The body plan and degree of specialization point to certain similarities between Omphreus (O.) bjelasicensis sp. n. and some subspecies of O. (O.) morio from Montenegro, viz., O. (O.) morio beckianus, O. (O.) morio morio, and O. (O.) morio malissorum. A more careful examination, however, reveals clear differences of character states between these four taxa. Thus, Omphreus (O.) bjelasicensis sp. n. is distinct from O. (O.) morio beckianus in body length (18.67 mm vs. 16.00– 18.00 mm), the antennae/body length ratio (0.57 vs. 0.55), presence/absence of the median labrum convexity (present vs. absent), the antennomere II/III length ratio (II = III vs. II < III), the head width/length ratio (0.935 vs. 1.03), the head/pronotum length ratio (0.64 vs. 0.61), the pronotum width/length ratio (0.87 vs. 0.92), shape of the posterior part of the lateral pronotal margins (strongly concave vs. weakly concave), and the elytral width/length ratio (elytra somewhat more than two thirds longer than wide vs. elytra about two thirds longer than wide). Omphreus (O.) bjelasicensis sp. n. differs from O. (O.) morio morio in body length (18.67 mm vs. 20.00 mm), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), shape of the anterior part of the lateral pronotal margins (less convex vs. more convex), and the elytral width/length ratio (elytra slightly more than two thirds longer than broad vs. elytra almost twice as long as broad). Further, O. (O.) bjelasicensis sp. n. differs from O. (O.) morio malissorum in body length (18.67 mm vs. 17.00–18.00 mm), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), and the elytral width/length ratio (elytra slightly more than two thirds longer than broad vs. elytra almost twice as long as broad). Omphreus (O.) bjelasicensis sp. n. also differs from Omphreus (O.) wohlberedti in body size (18.67 mm vs. 26.00 mm), size of the head (proportionally larger vs. proportionally smaller), form of the apical part of the first antennomere (sharply widening vs. gradually widening), the I/II–IV antennomere length ratio (I < II–IV vs. I = II–IV), the pronotum width/length ratio

(pronotum much longer than wide vs. pronotum somewhat longer than wide), shape of the anterior part of the pronotum (less widened and rounded vs. wider and more rounded), presence/absence of a row of points in the second interstrial spaces (absent vs. present), the elytral width/length ratio (elytra slightly more than two thirds longer than broad vs. elytra more than twice as long as broad), and the development of brush-like setation on the ventral side of the two fore male protarsomeres (well-developed vs. less developed). The diagnostic differences between Omphreus (O.) bjelasicensis sp. n. and Omphreus (O.) prekornicensis sp. n. are presented in the differential diagnosis of the species Omphreus (O.) prekornicensis sp. n. Bionomy and distribution. The new species inhabits high-altitude localities on Mt. Bjelasica in Biogradska Gora National Park, especially epigean and endogean habitats. These beetles are soil-dwellers, their nutrition being the same as in the species Omphreus (O.) prekornicensis sp. n. Like it, the given species represents an endemic relict form, probably of Tertiary origin. Acknowledgements We are especially indebted to Prof. Dr. I. Lichtscheidl and Mr. G. Eder (Institute of Cell Imaging and Ultrastructure Research, Faculty of Life Sciences, University of Vienna) for their valuable help in taking digital photos of the studied beetles. This study was financially supported by the Serbian Ministry of Science (Grant No. 143053).

References Bousquet Y. 2003. Tribe Omphreini Ganglbauer, 1891, pp. 445– 446. In: L¨ obl I. & Smetana A. (eds), Catalogue of Palaearctic Coleoptera, Vol. 1, Archostemata – Myxophaga – Adephaga, Apollo Books, Stenstrup, Denmark. Drovenik B. & Peks H. 1999. Coleoptera. Catalogus Faunae. Carabiden der Balkanl¨ ander. Coleoptera Carabidae. Schwanfelder Coleopterologische Mitteilungen (Neuauflage) 1: 1– 123. Trautner J. & Geigenm¨ uller K. 1987. Sandlaufk¨ afer. Laufk¨ afer. Illustrierter Schl¨ ussel zu den Cicindeliden und Carabiden Europas. Verlag Josef Margraf, Aichtal, 488 pp. Winkler A. 1933. Revision der Omphreus-Arten (Carab.). Koleopterologische Rundschau 19 (3–4): 115–120. Received January 9, 2007 Accepted February 15, 2008