Carabus (Tribax) satyrus Kurnakov 1962

Large form, males 31.1–35.7 (33.8) mm, females 32.2–39.8 (36.1) mm long, slender. Underside black, dorsum normally with bright metallic lustre, green, dark blue, bronze, crimson, violet, seldom black, with transitional color forms in males, females less nitidous, sometimes matte. Body shape is the narrowest within the studied species complex. Head not inflated. Pronotum extremely variable in shape, from subquadrate to transverse, more often cordate, lateral sides with deep hollow before hind angles. PW/PL = 1.13–1.43 (1.32) in males and 1.03–1.57 (1.37) in females, hind angles strongly protruding backwards and sidewards, pointed apically. Median groove smooth to completely obliterated in some female specimens. Disk transversally rugose, with rugosity gradually strengthening towards median groove. Elytra strongly elongate to ovate (generally the most slender within studied species complex), more elongate in males, with inconspicuous depression in the middle in some female specimens. EL/EW = 1.64–2.02 (1.84) in males and 1.60–1.97 (1.80) in females. Elytral sculpture nearly identical in males and females, forming precise series of elongate links. The main morphometric measurements of the studied populations are presented in Table 2. Morphometric characteristic of males (n = 67) and females (n = 68) of Carabus (Archiplectes) pseudopshuensis. Species/subspecies, locality, number of specimens studied GBL* SBL HW PW PB PL EL EW PW/PL PW/PB EL/EW EL/PL EW/PW Males pseudopshuensis vall. riv. Belaya, 1000 m, 1 ex 30.0 27.0 4.5 7.0 5.0 4.5 17.0 9.0 1.56 1.4 1.89 3.78 1.29 pseudopshuensis vall. riv. Byaul, 1100 m, 5 ex 29.0–32.0 30.6 26.0–29.0 27.6 4.5–5.5 4.84 6.5–7.5 6.84 5.5–6.0 5.74 5.0 16.5–18 17.36 9.0–10.0 9.56 1.3–1.5 1.37 1.17–1.25 1.19 1.78–1.86 1.82 3.3–3.6 3.47 1.33–1.49 1.4 pseudopshuensis Village Sanchara, 800 m, 5 ex 28.4–32.75 30.11 26.2–30.3 27.77 4.4–5.45 4.85 6.6–8.35 7.18 5.0–6.1 5.51 4.75–5.4 5.18 16–18.85 17.24 9.15–11.1 10.14 1.28–1.55 1.39 1.26–1.37 1.3 1.64–1.84 1.7 3.19–3.49 3.33 1.33–1.49 1.42 pseudopshuensis Village Bitaga (Pskhu), 700 m, 20 ex 26.45–32 29.47 24.25–30.0 27.22 4.25–5.0 4.69 5.9–7.2 6.7 4.4–5.8 5.24 4.5–5.5 5.09 14.75–18 16.7 8.8–11 9.87 1.2–1.41 1.32 1.09–1.4 1.28 1.51–1.86 1.69 2.91–3.5 3.28 1.34–1.66 1.48 pseudopshuensis vall. riv. Reshevie, 700 m, 20 ex 29.0–32.0 30.39 26.55–30.0 28.22 4.4–6.0 5.13 6.35–8.2 7.17 4.85–7.0 5.75 4.9–5.5 5.16 16.5–22.7 17.84 9.2–12.4 10.6 1.21–1.64 1.39 1–1.41 1.25 1.45–2.17 1.69 3.09–4.63 3.47 1.33–1.6 1.48 pseudopshuensis Dou Pass, 1300 m, 1 ex 31.0 29.0 5.0 7.0 6.0 6.0 17.0 10.4 1.17 1.17 1.63 2.83 1.49 pseudopshuensis vall. riv. Bzyb, 900 m, 5 ex 29.0–31.0 30.3 27.0–29.5 28.4 5.0 6.5–7.0 6.86 5.0–6.5 5.84 5.0–6.0 5.46 17.0–18.0 17.56 10.0–11.0 10.32 1.17–1.4 1.26 1.08–1.3 1.18 1.59–1.8 1.7 3–3.56 3.23 1.43–1.69 1.51 pseudopshuensis Village Serebryanyi, 600 m, 10 ex 28.0–31.0 29.4 26.0–29.0 27.45 4.8–6.0 5.28 6.5–8.0 7.21 5.0–7.0 6.11 5.2–5.7 5.43 16.0–18.0 17.5 9.6–11.6 10.58 1.18–1.51 1.33 1–1.33 1.19 1.38–1.88 1.66 2.91–3.4 3.22 1.37–1.55 1.47 Females pseudopshuensis vall. riv. Belaya, 1000 m, 1 ex 31.0 29.0 5.0 7.5 5.5 5.0 17.0 8.0 1.5 1.36 2.13 3.4 1.07 pseudopshuensis vall. riv. Byaul, 1100 m, 5 ex 31.0–33.0 32.26 28.5–31.0 30.1 4.5–5.5 5.1 7.0–8.0 7.4 5.0–7.8 6.16 4.5–5.5 5.0 15.3–19.0 16.96 9.0–11.0 10.08 1.4–1.56 1.48 1.03–1.4 1.23 1.47–1.89 1.69 3.06–3.78 3.4 1.29–1.43 1.36 pseudopshuensis Village Sanchara, 800 m, 4 ex 31.0–32.2 31.63 29.0–29.65 29.26 4.8–5.55 5.15 7.0–8.0 7.58 5.85–6.5 6.11 5.3–5.0 5.33 18.25–19.0 18.53 10.45–10.8 10.56 1.32–1.56 1.42 1.2–1.28 1.24 1.69–1.81 1.75 3.34–3.8 3.49 1.31–1.49 1.4 pseudopshuensis Village Bitaga (Pskhu), 700 m, 20 ex 29.55–33 31.99 27.75–30.5 29.46 4.65–5.35 5.01 7.0–8.5 7.42 5.35–6.5 5.98 5.0–5.8 5.4 16.8–19.4 17.98 9.6–10.95 10.25 1.21–1.55 1.38 1.08–1.42 1.24 1.6–1.98 1.76 3.08–3.59 3.33 1.13–1.51 1.39 pseudopshuensis vall. riv. Reshevie, 700 m, 20 ex 29.0–34.0 31.58 27.0–32.0 29.13 4.5–5.45 4.92 6.5–8.55 7.4 5.2–6.8 6.04 4.75–6.3 5.25 16.5–19.4 18.19 9.0–11.25 10.4 1.25–1.63 1.42 1.1–1.35 1.23 1.59–1.9 1.75 3.02–3.8 3.48 1.3–1.62 1.41 pseudopshuensis Dou Pass, 1300 m, 3 ex 31.5–33.0 32.17 29.5–31.0 30.17 5.0 7.3–7.8 7.53 6.0–6.3 6.1 5.0–5.3 5.17 17.5–19.0 18 9.6–11.6 10.53 1.38–1.5 1.46 1.22–1.25 1.23 1.64–1.82 1.71 3.5–3.58 3.48 1.28–1.59 1.4 pseudopshuensis vall. riv. Bzyb, 900 m, 5 ex 30.0–33.0 31.5 27.5–30.5 29.2 5.2–5.5 5.4 7.0–7.5 7.14 6.0 5.0–6.0 5.36 17.0–18.0 17.7 10.0–12.0 11.08 1.25–1.4 1.34 1.17–1.25 1.19 1.5–1.7 1.6 3.0–3.6 3.32 1.43–1.63 1.55 pseudopshuensis Village Serebryanyi, 600 m, 10 ex 28.5–32.0 30.3 26.5–30.0 28.3 4.5–5.8 5.22 5.8–8.0 6.8 5.0–7.0 5.9 5.0–6.5 5.58 16.5–19.5 17.75 9.6–11.0 10.32 0.89–1.44 1.23 1.07–1.2 1.16 1.63–1.84 1.72 2.83–3.45 3.19 1.28–1.83 1.53 for abbreviations see text. Morphometric characteristic of males (n = 16) and females (n = 31) of Carabus (Archiplectes) satyrus and males (n = 83) and females (n = 91) of Carabus (Archiplectes) besleticus subspecies. Species/subspecies, locality, number of specimens studied GBL* SBL HW PW PB PL EL EW PW/PL PW/PB EL/EW EL/PL EW/PW Males satyrus Village Merkheul, 120–230 m, 16 ex. 31.1–35.7 33.84 28.6–33.1 31.22 5.0–5.65 5.39 6.8–8.6 7.84 5.75–6.85 6.33 5.65–6.6 5.97 18.2–21.4 19.75 9.95–11.35 10.73 1.13–1.43 1.32 1.06–1.36 1.24 1.64–2.02 1.84 3.03–3.68 3.32 1.22–1.49 1.37 besleticus resheviensis vall. riv. Reshevie, 700 m, 29 ex. 34.5–39.0 37.62 32.6–37.0 35.15 5.75–6.35 6.03 8.0–9.5 8.58 6.8–8.2 7.19 5.5–7.0 6.27 21–22.7 21.45 12.1–15 13.51 1.18–1.55 1.37 1.1–1.3 1.19 1.42–1.78 1.59 3–3.79 3.43 1.41–1.83 1.58 besleticus adzinbai Mt. Akibakhu, 2000 m, 1 ex. 31.0 29.0 6.5 7.5 6.5 6.0 20.0 12.4 1.25 1.15 1.61 3.33 1.65 besleticus mtsaranus vall. riv. Mtsara, 550–750 m, 10 ex. 35.25–37.1 35.77 31.8–34.0 32.82 5.5–5.75 5.59 7.5–8.85 8.19 6.4–7.2 6.81 5.55–6.9 6.1 19.6–22.15 20.79 11.15–12.55 11.87 1.14–1.48 1.35 1.15–1.26 1.2 1.65–1.85 1.75 2.84–3.87 3.42 1.3–1.63 1.45 besleticus besleticus Mt. Birtzkha, 300 m, 20 ex. 32.0–38.0 34.68 30.5–34.65 32.29 5.4–6.3 5.87 7.55–9.75 8.36 6.25–8.0 7.21 5.65–6.5 6.12 20–22.5 21.34 10.9–13 12.23 1.26–1.53 1.37 1.03–1.33 1.17 1.62–1.86 1.75 3.29–3.75 3.49 1.23–1.63 1.47 besleticus dsykhvensis Mt. Dzykhva, 2000 m, 19 ex. 23.3–26.5 25.2 21.7–25.4 23.41 4.0–5.0 4.46 5.65–6.4 6.13 4.6–5.6 5.06 4.0–4.6 4.29 13.4–15.8 14.56 8.0–9.2 8.65 1.2–1.58 1.43 1.07–1.37 1.22 1.56–1.85 1.68 3.08–3.59 3.39 1.29–1.51 1.41 besleticus duripshensis Bzybian Mt. Range. 420–550 m, 2 ex. 32.45–32.5 32.48 29.75–30.0 29.88 4.75–5.3 5.03 7.2–7.6 7.4 5.65–5.75 5.7 5.15–5.85 5.5 19.1 19.1 10.6–10.8 10.7 1.30–1.40 1.35 1.27–1.32 1.30 1.77–1.80 1.79 3.26–3.71 3.49 1.42–1.47 1.45 besleticus napraensis NW of Bzybian Mt. Range, 1900–2000 m, 2 ex. 28.8–30.7 29.75 26.45–28.15 27.3 4.85–5.15 5.0 6.65–6.85 6.75 5.25–5.8 5.53 5.25–5.25 5.25 17.0–17.65 17.33 9.8–10.55 10.18 1.27–1.31 1.29 1.18–1.27 1.22 1.67–1.74 1.70 3.24–3.36 3.30 1.48–1.54 1.51 Females satyrus Village Merkheul, 120–230 m, 31 ex. 32.2–39.75 36.11 29.5–37.25 33.24 5.25–6.3 5.81 7.5–9.5 8.58 6.0–7.6 6.9 5.35–8.0 6.33 18.0–24.15 21.04 10.9–13.4 11.71 1.03–1.57 1.37 1.08–1.36 1.24 1.6–1.97 1.80 2.48–3.79 3.35 1.22–1.51 1.37 besleticus resheviensis vall. riv. Reshevie, 700 m, 28 ex 37.15–43 40.61 34.15–41 37.9 5.7–7.2 6.6 8.25–10 9.43 6.5–9.0 7.92 5.6–7.2 6.5 19.8–25 23.59 12.25–16 13.74 1.29–1.67 1.45 1.06–1.33 1.2 1.52–1.92 1.72 2.96–4.08 3.64 1.3–1.67 1.46 besleticus adzinbai Mt. Akibakhu, 2000 m, 10 ex. 32–35.75 33.33 30.0–32.9 31.24 5.85–6.5 6.06 7.5–9.5 8.33 6.5–8.5 7.31 6.0–7.5 6.66 20.65–22.5 21.7 11.25–13 12.65 1.14–1.42 1.25 1.0–1.26 1.14 1.64–1.84 1.72 2.93–3.67 3.27 1.32–1.73 1.53 besleticus mtsaranus vall. riv. Mtsara, 550–750 m, 10 ex. 37.5–41.25 39.28 34.5–37.75 36.11 5.8–6.25 6.01 8.3–9.5 9.12 7.0–8.15 7.43 5.85–7.0 6.51 21.7–24 22.74 12.1–13.5 12.67 1.26–1.56 1.41 1.13–1.32 1.23 1.73–1.98 1.8 3.29–3.86 3.5 1.29–1.47 1.39 besleticus besleticus Mt. Birtzkha, 300 m, 20 ex. 38.0–40.75 38.95 35.15–38.0 36.45 5.85–6.5 6.27 8.5–11.0 9.74 7.0–9.0 8.19 6.0–7.2 6.68 22.45–24.0 23.22 12.15–14.4 13.06 1.25–1.62 1.46 1.11–1.31 1.19 1.64–1.86 1.78 3.19–3.88 3.48 1.13–1.47 1.35 besleticus besleticus Village Abzhakva, 70 m, 1 ex. 39.0 36.0 6.5 10.3 8.1 7.0 22.5 13.5 1.46 1.27 1.67 3.21 1.32 besleticus dsykhvensis Mt. Dzykhva, 2000 m, 18 ex. 23.3–26.5 25.2 23.65–26.3 25.18 4.0–4.9 4.44 6.0–7.3 6.47 5.0–6.8 5.53 4.2–4.9 4.44 14.65–17.0 15.5 8.65–10 9.38 1.3–1.62 1.46 1.03–1.3 1.17 1.5–1.79 1.66 3.33–3.67 3.49 1.34–1.59 1.45 besleticus duripshensis Bzybian Mt. Range, 420–550 m, 2 ex. 32.5–32.6 32.55 30.0–30.3 30.15 5.25–5.25 5.25 7.25–7.3 7.28 5.75–6.3 6.03 5.65–5.75 5.7 19.0–20.0 19.5 10.5–10.75 10.63 1.27–1.28 1.28 1.16–1.26 1.21 1.81–1.86 1.83 3.36–3.48 3.42 1.45–1.47 1.46 besleticus napraensis NW of Bzybian Mt. Range, 1900–2000 m, 2 ex. 31.7–33.7 32.7 29.35–30.6 29.98 5.35–5.5 5.43 7.6–7.9 7.75 6.35–6.45 6.4 5.15–5.3 5.23 19.0–20.35 19.68 10.5–12.0 11.25 1.48–1.49 1.48 1.20–1.22 1.21 1.70–1.81 1.75 3.69–3.84 3.76 1.38–1.52 1.45 for abbreviations see text. Lamella of aedeagus short, of triangular form. Structure of endophallus testifies to the assignment of this form to the reitteri-group (since the previous authors did not properly analyze this morphological structure, it lead them to an erroneous interpretation of the subspecific structure of this species). Saccellus short, of rectangular shape, resembling saccellus of Carabus (Archiplectes) juenthneri, with S-shaped fold on the right, paracellar lobe rounded, protruding laterally, endophallus with asymmetric position of inner structures (when viewed frontally). Ostium lobe small, sometimes faintly developed. Aggonoporius small.

Populates low mountain forest belts in an interfluve of the r. Kodor and Kelasur, from altitude of 50 m up to 500 m.