Holotype male labeled as follows: “eastern Bosnia and Herzegovina, southern slope of Mt. Zelengora, Čemerno, 1,450 m a.s.l., near Gacko, 22.VI–14.VII.2012, from pitfall traps, leg. F. Sandel” (white label, printed) / Holotypus Omphreus (Omphreus) morio sandeli ssp. n. S. Ćurčić & R. Sciaky det. 2014” (red label, printed) (IZFB). Paratypes: four males and one female, same data as for holotype (CRS); one male labeled as follows: “eastern Bosnia and Herzegovina, southern slope of Mt. Zelengora, Čemerno, 1,600 m a.s.l., near Foča, 07–24.VIII.2011, from pitfall traps, leg. F. Sandel” (CRS); four males and two females labeled as follows: “eastern Bosnia and Herzegovina, Mt. Maglić, Tjentište, 1,450 m a.s.l., near Foča, 23.VI–13.VII.2012, from pitfall traps, leg. F. Sandel” (CRS); two males and one female labeled as follows: “eastern Bosnia and Herzegovina, northern slope of Mt. Zelengora, Tjentište, 1,450 m a.s.l., near Foča, 23.VI–13.VII.2012, from pitfall traps, leg. F. Sandel” (CRS). All paratypes are labeled with white, printed locality labels and with red printed labels “Paratypus Omphreus (Omphreus) morio sandeli ssp. n. S. Ćurčić & R. Sciaky det. 2014”.

Size large: TL: R 16.88–19.73 mm (M 18.49 mm). Body elongate; elytra ovoid (Figure 4). Body color black, mouthparts, apical antennomeres, and tarsi black-brownish. Tegument shiny, except slightly matt elytra.

Figures 4–8. 

Omphreus (Omphreus) morio sandeli ssp. n. from eastern Bosnia and Herzegovina, southern slope of Mt. Zelengora, Čemerno, 1,450 m a.s.l., near Gacko. 4 holotype male, habitus (dorsal view) 5 holotype male, aedeagus (dorsal view) 6 holotype male, aedeagus (lateral view) 7 holotype male, abdominal sternite IX (urite) 8 paratype female, genitalia. Scale bars 5 mm (4) and 1 mm (5–8).

Head rounded, somewhat elongated [HW/HL: M 0.96 (R 0.91–1.05)], narrower (HW/PW: M 0.69) than pronotum (Figure 4). Head below eye level somewhat constricted. Labrum broad, medially rounded, carrying four setae. Epistome large, concave anteriorly, with two setae. Both vertex and occiput wrinkled. Frontal foveae deepened and long. Gula bisetose. Mandibles elongated, sickle-shaped, broadened basally. Labial palpomere 1 short, without setae. The labial palpomeres 2 and 3 longer. Both labial and maxillar palpomere 3 broadened distally and densely pubescent. AL: M 10.41 mm (R 9.25–11.16 mm). Antennae pubescent from antennomere 4. AL/TL: M 0.56 (R 0.51–0.58). 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.

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.

Elytra ovoid, relatively wide, rounded laterally, EW/EL: M 0.58 (R 0.54–0.61) (Figure 4). EL/EW: M 1.73. EW: M 6.26 mm in males, while 6.36 mm in females. Elytral striae shallow, weakly punctate, points somewhat deeper basally. Elytral intervals flattened. Scutellum large, triangular. Scutellar stria present, but without scutellar puncture. Elytra widest around at the middle. A setiferous puncture in interval 7 basally (close to stria 6), 4–6 setiferous punctures in interval 7 medially (close to stria 7), and two setiferous punctures situated on stria 7 apically on each elytron. The punctures deep and large. Shoulders rounded. Umbilicate series regular, with the setae densely distributed. Elytral disc somewhat convex distally.

Protarsomeres 1 and 2 widened in males. Metacoxae long and rounded. Tarsal claws elongate, glabrous, smooth (Figure 4).

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).

Male abdominal sternite IX (urite) large, sub-triangular (Figure 7).

Both gonocoxites and gonosubcoxites IX as presented in Figure 8. Gonocoxites IX wide and elongated, somewhat curved, gradually narrowing distally, each with a rounded apex, basally joined with massive rounded gonosubcoxites IX.

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.

The new subspecies is compared here with the morphologically nearest subspecies of Omphreus (Omphreus) morio Dejean, 1828. These are O. (O.) morio beckianus Ganglbauer, 1888, O. (O.) morio serbicus Winkler, 1933, and O. (O.) morio durmitorensis ssp. n. (the first antennomere being club-like distally, somewhat shorter than the following three antennomeres combined in all the subspecies mentioned) (Ganglbauer 1888, Winkler 1933).

Omphreus (O.) morio sandeli ssp. n. differs from O. (O.) morio beckianus in the shape of the elytra (distinctly rounded laterally vs. slightly rounded laterally), the elytra length/width ratio (M 1.73 vs. around 1.66), and the body length (R 16.88–19.73 mm vs. 16–18 mm) (Ganglbauer 1888, Winkler 1933). The former taxon is distributed both on Mt. Zelengora and Mt. Maglić, while the latter one inhabits Mt. Visočica (the type locality) (Ganglbauer 1888). In the literature Omphreus (O.) morio beckianus is reported to have a wide distribution in Bosnia and Herzegovina, Montenegro and Serbia (from central and southern Bosnia to border with Montenegro, southwestern Serbia and northern Herzegovina, including the high-altitude area of the surroundings of Sarajevo, Mts. Ivan, Bjelašnica, Volujak and Durmitor and the Lim River Valley) (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 (O.) morio sandeli ssp. n. differs from O. (O.) morio serbicus in the shape of the elytra (elongate vs. short), the elytra length/width ratio (M 1.73 vs. around 1.50), and the body length (R 16.88–19.73 mm vs. 15–16 mm) (Winkler 1933). Omphreus (O.) morio sandeli ssp. n. is found in eastern Bosnia and Herzegovina (Mt. Zelengora and Mt. Maglić), while O. (O.) morio serbicus lives in southwestern Serbia (Mt. Murtenica and Mt. Zlatibor) (Winkler 1933).

Eventually, Omphreus (O.) morio sandeli ssp. n. differs from O. (O.) morio durmitorensis ssp. n. in the antennal length (M 10.41 mm vs. 10.38 mm), 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).

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.

So far known only from the type locality (Mt. Zelengora) and the nearby Mt. Maglić, eastern Bosnia and Herzegovina.

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.

Holotype male labeled as follows: “northwestern Montenegro, Mt. Durmitor, Sedlo pass, 2,100–2,200 m a.s.l., near Žabljak, 28.VI–17.VII.2014, from pitfall traps, leg. S. Ćurčić” (white label, printed) / Holotypus Omphreus (Omphreus) morio durmitorensis ssp. n. S. Ćurčić & R. Sciaky det. 2014” (red label, printed) (IZFB). Paratypes: one female, same data as for holotype (IZFB); one male labeled as follows: “northwestern Montenegro, Mt. Durmitor, Sedlo pass, 2,100 m a.s.l., 28–29.VI.2014, from pitfall traps, leg. S. Ćurčić & D. Antić” (IZFB); one male labeled as follows: “northwestern Montenegro, Mt. Durmitor, Žabljak, 1,950 m a.s.l., VIII.2006, leg. P. Zanandrea” (CRS); one female labeled as follows: “northwestern Montenegro, Mt. Durmitor, Sedlo pass, 2,200 m a.s.l., 07.VIII.2007, leg. F. Sandel” (CRS). All paratypes are labeled with white, printed locality labels and with red printed labels “Paratypus Omphreus (Omphreus) morio durmitorensis ssp. n. S. Ćurčić & R. Sciaky det. 2014”.

Size large: TL: R 17.73–18.91 mm (M 18.13 mm). Body elongate; elytra ovate (Figure 9). Body color black, mouthparts, apical antennomeres, and tarsi black-brownish. Tegument shiny, except slightly matt elytra.

Figures 9–13. 

Omphreus (Omphreus) morio durmitorensis ssp. n. from northwestern Montenegro, Mt. Durmitor, Sedlo pass, 2,100–2,200 m a.s.l., near Žabljak. 9 holotype male, habitus (dorsal view) 10 holotype male, aedeagus (dorsal view) 11 holotype male, aedeagus (lateral view) 12 holotype male, abdominal sternite IX (urite) 13 paratype female, genitalia. Scale bars 5 mm (9) and 1 mm (10–13).

Head rounded, somewhat elongated [HW/HL: M 0.95 (R 0.93–0.99)], shorter and narrower (HW/PW: M 0.695) than pronotum (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.

Pronotum sub-campaniform, elongate, PW/PL: M 0.91 (R 0.90–0.92). Fore angles somewhat prominent, rounded, hind angles obtuse, somewhat rounded (Figure 9). Lateral margins well developed, thickened, arcuate anteriorly, then sinuate and narrowing posteriorly. Anterior pronotal margin somewhat concave, while the base strongly concave. Pronotum widest in front of the fore 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 half of pronotum length. Pronotal disc somewhat convex proximally.

Elytra ovate, relatively long, arcuate laterally, EW/EL: M 0.56 (R 0.53–0.58) (Figure 9). EL/EW: M 1.79. EW: M 5.94 mm in males, 6.12 mm in females. Elytral striae shallow, weakly punctate, points somewhat deeper basally; elytral intervals flattened. Scutellum large, triangular; scutellar stria present, but without scutellar punctures. Elytra widest slightly below the middle; one setiferous puncture in interval 7 basally (close to stria 6), four or five setiferous punctures in interval 7 medially (close to stria 7), and two setiferous punctures situated on stria 7 apically on each elytron; the punctures deep and large. Shoulders obtusely rounded. Umbilicate series regular, with the setae densely distributed. Elytral disc somewhat convex distally.

Protarsomeres 1 and 2 widened in males. Metacoxae long and rounded. Tarsal claws elongated, glabrous, without teeth (Figure 9).

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).

Male abdominal sternite IX (urite) large, sub-triangular (Figure 12).

Both gonocoxites and gonosubcoxites IX as presented in Figure 13. Gonocoxites IX wide and elongated, somewhat curved, gradually narrowing distally, each with a pointed apex, basally joined with massive rounded gonosubcoxites IX.

The new subspecies is compared here with the morphologically nearest subspecies of Omphreus (Omphreus) morio. These are O. (O.) morio beckianus, O. (O.) morio serbicus, and O. (O.) morio sandeli ssp. n. In all these subspecies the first antennomere is club-like distally, somewhat shorter than the following three antennomeres combined (Ganglbauer 1888, Winkler 1933).

Omphreus (O.) morio durmitorensis ssp. n. differs from O. (O.) morio beckianus in the shape of the elytra (arcuate laterally vs. somewhat rounded laterally), the elytra length/width ratio (M 1.79 vs. around 1.66), and the body length (R 17.73–18.91 mm vs. 16–18 mm) (Ganglbauer 1888, Winkler 1933). The former taxon is distributed on Mt. Durmitor (northwestern Montenegro), while the latter inhabits Mt. Visočica (central Bosnia and Herzegovina) (the type locality) (Ganglbauer 1888). Apfelbeck (1904) and Drovenik (1984) recorded the presence of an Omphreus on Mt. Durmitor reported as O. (O.) morio beckianus. While the former author recorded it for the mountain without precise localities, the latter author found it in several different sites (Crno Jezero Lake surroundings, forests below Mali Štulac, Zminje Jezero Lake surroundings, Veliki Štulac, Ališnica, Međed, Velika Karlica, Savin Kuk, Lokvice, Bobotov Kuk, Dobri Do, Sedlo, Todorov Do, Prutaš, Škrčka Jezera Lakes surroundings, Bolj, forest below Ćurevac, Nagorje, and Šljeme) (Apfelbeck 1904, Drovenik 1984). On the basis of the material collected from Mt. Durmitor (at Sedlo and Žabljak sites) loaned by a few colleagues, as well as the one collected by the first and third author (S.Ć. and D.A.), we have concluded that the taxon from Mt. Durmitor actually represents a new subspecies, O. (O.) morio durmitorensis ssp. n.

Omphreus (O.) morio durmitorensis ssp. n. differs from O. (O.) morio serbicus in the shape of the elytra (elongate, arcuate laterally vs. short, rounded laterally), the elytra length/width ratio (M 1.79 vs. around 1.50), and the body length (R 17.73–18.91 mm vs. 15–16 mm) (Winkler 1933). Furthermore, they inhabit quite distant areas: Omphreus (O.) morio durmitorensis ssp. n. is found in northwestern Montenegro (Mt. Durmitor), while O. (O.) morio serbicus lives in southwestern Serbia (Mt. Murtenica and Mt. Zlatibor) (Winkler 1933).

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.

The new subspecies is named after Mt. Durmitor, its terra typica.

So far known only from the type locality, Mt. Durmitor, northwestern Montenegro.

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.