The genus Omphreus in Bosnia and Herzegovina and Montenegro, with two new subspecies of O. morio (Coleoptera, Carabidae, Omphreini)

Abstract Two new ground beetle subspecies, Omphreus (Omphreus) morio sandeli ssp. n. (from Mts. Zelengora and Maglić, eastern Bosnia and Herzegovina) and Omphreus (Omphreus) morio durmitorensis ssp. n. (from Mt. Durmitor, northwestern Montenegro) are here described and diagnosed. The male and female genitalia and other taxonomically important characters are illustrated. The new taxa are distinctly different from the nearest relatives and represent both endemics and relicts inhabiting limited high-altitude Dinaric areas in Bosnia and Herzegovina and Montenegro. A key to Omphreus taxa from Montenegro and a key to subspecies of Omphreus (Omphreus) morio Dejean, 1828 are presented.

These were studied in the laboratory of the Institute of Zoology, University of Belgrade -Faculty of Biology, Belgrade, Serbia. The beetle specimens were dissected, studied, and imaged. Dry specimens and the genitalia were glued onto separate rectangular paper labels situated on the same pin.
Carl Zeiss -Stemi 2000 and Carl Zeiss Discovery V8 binocular stereomicroscopes with a Canon G10 digital camera, as well as Nikon Eclipse E100 microscope with a Moticam 2000 digital camera attached were used in the study.

M
mean value for certain measurements R range of the total measurements performed TL total body length (measured from the anterior margin of clypeus to the apex of elytra) HW/HL ratio maximum width of head/length of head HW/PW ratio maximum width of head/maximum width of pronotum, as greatest transverse distance AL total antennal length including the scape AL/TL ratio total antennal length including the scape/total body length (measured from the anterior margin of clypeus to the apex of elytra) PW/PL ratio maximum width of pronotum, as greatest transverse distance/length of pronotum (measured along the median line) EW/EL ratio maximum width of elytra/length of elytra (as linear distance measured along the suture from the elytral base to the apex) EL/EW ratio length of elytra (as linear distance measured along the suture from the elytral base to the apex)/maximum width of elytra EW maximum width of elytra  Figure 4). Body color black, mouthparts, apical antennomeres, and tarsi black-brownish. Tegument shiny, except slightly matt elytra.
Pronotum sub-campaniform, elongate, PW/PL: M 0.91 (R 0.885-0.94). Fore angles somewhat prominent, rounded, hind angles obtuse, well rounded ( Figure 4). Lateral margins well developed, thickened, arcuate anteriorly, then sinuate and narrowing posteriorly. Anterior pronotal margin somewhat concave, base strongly concave. Pronotum widest between its fore fourth and third. Lateral furrows narrow and shallow, with four anterior setae, one median seta, and one posterior seta each. Median furrow long and deep. Basal foveae deep and long, slightly shorter than one half of pronotum length. Pronotal disc somewhat convex proximally.
Aedeagus long, median lobe somewhat widened sub-apically in dorsal view ( Figure  5), while curved and strongly widened sub-apically in lateral view ( Figure 6), with a straight long acute triangular apex ( Figures 5 and 6). Parameres wide, the right being much broader ( Figure 5). Basal bulb wide and short ( Figures 5 and 6).
Variability. It was noticed that the specimens from the population from the southern slope of Mt. Zelengora are of somewhat larger size and more elongate elytra (EW/EL: M 0.56) compared with the specimens belonging to the populations both from Mt. Maglić and the northern slope of Mt. Zelengora (EW/EL: M 0.59). Other characteristics, including the structure of aedeagus, do not show any significant differences among the three analyzed populations of the taxon.
Differential diagnosis and remarks.  (Apfelbeck 1894, 1904, Wohlberedt-Triebes 1909, Drovenik 1984). This appears to us very doubtful since Omphreus taxa (species or subspecies) mostly inhabit single mountain ranges (Winkler 1933). The specimens from Mt. Durmitor belong to a new subspecies described herein -Omphreus (O.) morio durmitorensis ssp. n.; Mt. Volujak is relatively close to Mt. Zelengora and Mt. Maglić and there is a possibility that the population from this mountain might actually belong to Omphreus (O.) morio sandeli ssp. n., but this has to be proven. The specimens from all known sites should be compared in the future (including the structure of the male genitalia) in order to conclude whether they belong to a single taxon (subspecies) or to different taxa. Additionally, a detailed study of all Omphreus (O.) morio subspecies (with comparisons of numerous morphological characteristics) would be needed in order to define their real taxonomic status (subspecies or species).
Omphreus ( , the shape of the hind pronotal angles (more rounded vs. less rounded), maximum width of the pronotum (between its fore fourth and third vs. in front of the fore third), the shape of the elytra (somewhat widened vs. more elongate), the elytral width/length ratio (M 0.58 vs. 0.56), the elytra length/width ratio (M 1.73 vs. 1.79), the elytral width (M 6.26 mm in males, 6.36 mm in females vs. 5.94 mm in males, 6.12 mm in females), maximum width of the elytra (around at the middle vs. slightly below the middle), the shape of the shoulders (rounded vs. obtusely rounded), the form of the median lobe (somewhat widened sub-apically in dorsal view, while curved and strongly widened sub-apically in lateral view, with a straight long acute triangular apex vs. strongly widened sub-apically in dorsal view, while arcuate, moderately widened and with a shallow concavity in the sub-apical part in lateral view, with a straight short rounded triangular apex), the shape of the basal bulb (wide and short vs. narrow and elongated), the shape of the male abdominal sternite IX (urite) (less elongate vs. more elongate), the form of the apex of the gonocoxites IX (rounded vs. pointed), and the total body length (M 18.49 mm vs. 18.13 mm).
Etymology. This new subspecies is named after Franco Sandel, friend of the second author and excellent collector, who collected the whole type series of this new subspecies allowing us to freely study the material.
Distribution. So far known only from the type locality (Mt. Zelengora) and the nearby Mt. Maglić, eastern Bosnia and Herzegovina.
Habitat. The subspecies prefers high-altitude habitats (1,450-1,600 m a.s.l.) on Mt. Zelengora (at both the northern and southern slopes) and Mt. Maglić in eastern Bosnia and Herzegovina. Type series of the new subspecies was collected by pitfall traps filled with alcoholic vinegar, placed at different sites in beech forests by the alpine meadows on both mountains, and on a pass and near the first hairpin bend after the pass on the southern slope of Mt. Zelengora.  (Figure 9). Head beyond eye level somewhat constricted. Labrum broad, medially rounded, carrying four setae. Epistome huge, concave anteriorly, with two setae. Both vertex and occiput wrinkled. Frontal foveae deepened and long. Gula bisetose. Mandibles elongated, sickle-formed, broadened basally. Labial palpomere 1 short, without setae. Labial palpomeres 2 and 3 longer. Both the labial palpomere 3 and maxillar palpomere 3 broadened distally and densely pubescent. AL: M 10.38 mm (R 10.07-10.64 mm). Antennae pubescent from antennomere 4. AL/TL: M 0.57 (R 0.54-0.60). Antennomere 1 club-like, sharply widened distally, with a few long setae distally, somewhat shorter than the following three antennomeres combined. Antennomere 2 slightly shorter than antennomere 3.
Aedeagus long, median lobe strongly widened sub-apically in dorsal view ( Figure  10), while arcuate, moderately widened and with a shallow concavity in the sub-apical part in lateral view (Figure 11), with a straight short rounded triangular apex ( Figures  10 and 11). Parameres wide, the right being much huger ( Figure 10). Basal bulb narrow and elongated (Figures 10 and 11).
The diagnostic differences between Omphreus (O.) morio durmitorensis ssp. n. and O. (O.) morio sandeli ssp. n. are presented in the differential diagnosis of the latter.
Etymology. The new subspecies is named after Mt. Durmitor, its terra typica. Distribution. So far known only from the type locality, Mt. Durmitor, northwestern Montenegro.
Habitat. The subspecies prefers high-altitude habitats (1,950-2,200 m a.s.l.) on Mt. Durmitor in northwestern Montenegro. The type series of the new subspecies was collected by pitfall traps filled with alcoholic vinegar, placed at different sites on the border between alpine meadows and rocks, up to a timberline on Mt. Durmitor.

Key to Omphreus taxa from Montenegro (modified after Winkler 1933 and Ćurčić et al. 2008b)
1 The The first two male protarsomeres neither broadened, nor brush-like setose on the underside. The second male protarsomere elongate. Seventh intervals of the matt elytra with 5-7 setiferous punctures each. Head behind eyes clearly narrowed. The first antennomere equally broadened towards the top, as long as the following three antennomeres combined (subgenus Neomphreus Winkler, 1933). Elytra elongately oval, laterally less rounded, twice as long as broad, with strongly impressed striae. Pronotum weakly heart-shaped, laterally very slightly rounded, for one fourth longer than broad.