Polydesmidae, Leach, 1815

Golovatch, Sergei, Evsyukov, Aleksandr & Reip, Hans, 2016, The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida), Zootaxa 4085 (1), pp. 1-51 : 47-48

Key to Polydesmidae species presently known to occur in the Caucasus

1. Adults with 20 segments, including telson. Collum often as broad as or broader than head. Gonopod telopodite often elongate and subgeniculate distally.................................................................... Polydesmus, 2

- Adults with 19 segments, including telson. Head broader than collum. Gonopod telopodite stout and never subgeniculate distally................................................................................... Brachydesmus , 5

2. Live coloration usually light brown to brown ( Figs 3A , 5A ). Paraterga not upturned, subhorizontal to slightly declivous, dorsum slightly, but clearly convex in the middle ( Figs 3A, 3H – 3J , 5A ). Collum clearly narrower than head ( Figs 3A, 3H , 5A )...................................................................................................... 3

- Coloration largely red to pinkish. Paraterga very broad, held high, about level to a flat mid-dorsal region, a few first postcollum paraterga slightly, but clearly upturned ( Figs 2E , 3B )........................................................ 4

3. Male prefemora strongly bulging laterad ( Fig. 5B ). Gonopod telopodite stout ( Fig. 5C )......... Polydesmus mediterraneus

- Male prefemora only slightly bulging laterad. Gonopod telopodite strongly elongate, subgeniculate distally ( Fig. 4C, 4D )............................................................................................ P. muralewiczi

4. Length ca 10 – 13 mm (male, female), width of midbody metazonae 1.25 – 1.7 (male) or 1.4 – 2.1 mm (female), metazonite to prozonite width ratio <2.0 ( Fig. 2 ). Collum about as broad as head ( Fig. 2A, 2B, 2F )......................... P. lignaui

- Length ca 13 – 19 mm (male, female), width of midbody metazonae 1.9 – 3.3 (male) or 2.4 – 3.5 mm (female), metazonite to prozonite width ratio> 2.0 ( Fig. 1 ). Collum clearly broader than head ( Fig. 1 ).............................. P. abchasius

5. Paraterga poorly-developed, their fore margins and anterolateral corners largely broadly and regularly rounded, not angular ( Fig. 10C ). Metatergal sculpture strongly obliterate ( Fig. 10A – 10D ). Gonopod exomere (ex) with a conspicuous densely pilose strip ( Fig. 10E – 10G )................................................................ Brachydesmus furcatus

- Paraterga variously developed, but their fore margins and anterolateral corners mostly clearly angulate (e.g. Figs 6 , 9A , 12 , 13B, 13E , 15B , 16B, 16E ). Metatergal sculpture often more strongly developed. Gonopod exomere without a pilose strip, but endomere often with densely pilose patches................................................................ 6

6. Tergal setae sharp ( Fig. 9A ). Gonopod ( Fig. 9B ) unipartite, slender, only slightly falcate, ventrally with several teeth or spines............................................................................................. B. superus

- Tergal setae mostly bacilliform or subclavate, only rarely sharpened. Gonopod telopodite bipartite, with a distinct exomere and a smaller, but evident endomere.

7. Adults <8 mm long and ≤ 1.0 mm wide. Gonopod exomere with only a single apical tooth ( Figs 28C – 28D , 29C – 29G ). Colchis, Black Sea coast area............................................................................ 8

- Adults ≥ 9 mm long and ≥ 1.0 mm wide.................................................................... 9

8. Length 6 – 7.5 mm, width of midbody metazonae 0.8 – 1.0 mm, metazonite to prozonite width ratio ca 1.5 – 1.6 ( Fig. 29A ). Male legs slightly, but clearly incrassate ( Fig. 29B ). Gonopod exomere longer ( Figs 29C – 29G ). Ajaria, Georgia.. B. simplex sp. n.

- Length 5 – 6 mm, width of midbody metazonae 0.65 – 0.7 mm, metazonite to prozonite width ratio ca 1.3 ( Fig. 28A ). Male legs slender ( Fig. 28B ). Gonopod exomere shorter ( Figs 28C – 28D ). Abkhazia and Sochi , Krasnodar Prov., Russia................................................................................................... B. kvavadzei sp. n.

9. Gonopod endomere (en) partially densely pilose and clearly bifid; exomere (ex) with 1 – 2 apical teeth e and i (the latter often subapical), often also with another subapical lobule or denticle l ( Figs 13G – 13H , 14C – 14E , 16G – 16H , 18D – 18E , 19D – 19E , 21D – 21F , 22 – 25 , 27B, 27D ). Extremely widespread (Map 2) and variable in all other characters, morphism outlined in Table 1...................................................................................... B. kalischewskyi

- Gonopods subfalcate, endomere (en) simple, more or less finger-shaped ( Figs 7 , 8E – 8F )........................... 10

10. Tergal setae mostly subclavate ( Fig. 6C – 6H ). Gonopod exomere (ex) more elaborate ( Fig. 7 ). Much of both Caucasus Major and Caucasus Minor, but not Hyrcania (Map 3)..................................................... B. assimilis

- Tergal setae sharp ( Fig. 9A ). Gonopod exomere (ex) simple ( Fig. 8F ). Hyrcania (Map 3)................... B. pigmentatus

Figures

FIGURE 1. Habitus of Polydesmus abchasius Attems, 1898. A: male from Gelgeluk, Abkhazia, lateral view. B: female from near Krasnaya Polyana, Krasnodar Prov., dorsal view. C: female ca 18 mm long from an unspecified locality in Abkhazia, dorsal view. Pictures A and B by K. Makarov, taken not to scale; watercolour C after Lignau (1915).

FIGURE 2. Habitus of Polydesmus lignaui Lohmander, 1936. A: male from Kherkhvashi, Svanetia, Georgia, dorsal view. B – E: anterior, middle and caudal parts of body, and entire body, respectively, of male from Sukhumi University Campus, Abkhazia, dorsal (B – D) and lateral views (E). F: male ca 12 mm long from an unspecified locality in Abkhazia, dorsal view. Pictures A – E by K. Makarov, taken not to scale; watercolour F after Lignau (1915).

FIGURE 3. Habitus of Polydesmus muralewiczi Lohmander, 1936 (A, H – J), P. lignaui Lohmander, 1936 (D – G) and P. abchasius Attems, 1898 (B, C). A, H – J: male from near Kursavka, Stavropol Prov., Russia, entire body, anterior body part, left half of metatergite 10 and caudal body part, respectively, dorsal views. B, C: male from Pasture Abago, Caucasian Nature Reserve, Adygea, Russia, anterior part of body, lateral view, and left half of metatergite 10, dorsal view, respectively. D – F: male from Dombai, Karachaevo-Cherkessia, Russia, anterior body part, left half of metatergite 10 and caudal part of body, respectively, dorsal views. G: female from Dombai, anterior part of body, lateral view. Picture A by K. Makarov, taken not to scale. Scale bar: 1.0 mm (B – J).

FIGURE 4. Left gonopods of Polydesmus abchasius Attems, 1898, P. lignaui Lohmander, 1936 and P. muralewiczi Lohmander, 1936 (A – C), and right gonopod of P. muralewiczi Lohmander, 1936 (D). A: male from near Lake Ritsa, Abkhazia, mesal view. B: male from Sukhumi Botanical Garden, Abkhazia, mesal view. C: male from near Kursavka, Stavropol Prov., Russia, mesal view. D: male lectotype from near Psebai, Krasnodar Prov., Russia (after Lohmander 1936). Designations: en, endomere; ex, exomere; d, exomere tooth; pu, hairy pulvillus; r, endomere rod. Scale bar: 0.2 mm (A – C), not to scale (D).

FIGURE 5. Polydesmus mediterraneus Daday, 1889: A: male from Gurzuf, Crimea, habitus, dorsal view. B – C: male from Yashtukha, Abkhazia, leg 7, lateral view, and left gonopod, mesal view, respectively. Designations: en, endomere, ex, exomere. Picture A by K. Makarov. Scale bars: 0.2 mm (B), 0.1 mm (C), not to scale (A).

FIGURE 6. Habitus of Brachydesmus assimilis Lohmander, 1936. A: male from Belakany, Azerbaijan. B – H: male (B), female (C – E) and male (F – H) from Upper Gunib, Dagestan, Russia. C, F: anterior part of body. D, G: right half of segment 10. E, H: caudal part of body. Pictures A and B by K. Makarov, taken not to scale. Line drawings C – H by S. Dashdamirov, scale bar: 1.0 mm.

FIGURE 7. Gonopods of Brachydesmus assimilis Lohmander, 1936. A – B: male from Mariamdjvari Nature Reserve, Georgia, lateral and mesal views, respectively. C: male from near Pass Krestovyi, Georgia. D: male syntype from Tbilisi, Georgia (after Lohmander 1936). Designations: en, endomere; ex, exomere; s, spine on exomere; t, tooth near base of exomere; k, spine at base of endomere; b, tooth at base of femorite. Scale bar: 0.1 mm (A – C), not to scale (D).

FIGURE 8. Brachydesmus pigmentatus Attems, 1951, males from Hyrcan Nature Reserve, Azerbaijan. A: habitus, entire body, dorsal view. B: anterior part of body, dorsal view. C: right half of metatergite 10, dorsal view. D: posterior part of body, dorsal view. E: gonopods in situ, ventral view. F: right gonopod, mesal view. Designations: en, endomere; ex, exomere; k, spine at base of endomere; t, tooth near base of femorite. Pictures A and E by K. Makarov, taken not to scale. Line drawings B – D by S. Dashdamirov, scale bar: 1.0 mm (B – D), 0.3 mm (F).

FIGURE 9. Brachydesmus superus Latzel, 1884. A: habitus, entire body, dorsal view, male from Stavropol, Russia. B: left gonopod, male from Zheleznovodsk, Stavropol Prov., Russia. Designation: pu, hairy pulvillus. Picture A taken by R. Zuev. Scale bars: 1.0 mm (A), 0.1 mm (B).

FIGURE 10. Brachydesmus furcatus Lohmander, 1936. A: habitus, entire body, dorsal view, male from Mount Kogot, Caucasian Nature Reserve, Krasnodar Prov., Russia. B – D: male from near Goryachiy Klyuch, Krasnodar Prov., Russia, B: anterior part of body, dorsal view; C: right half of metatergite 10, dorsal view; D: posterior part of body, dorsal view. E – F: male holotype of B. furcatus, right and left gonopods, lateral and mesal views, respectively (after Lohmander 1936); G: male holotype of B. furcatus exiguus Strasser, 1970, left gonopod, lateral view (after Strasser 1970). Designations: en, endomere; ex, exomere; t, tooth at base of femorite. Picture A by K. Makarov, taken not to scale. Line drawings B – D by S. Dashdamirov, scale bar: 1.0 mm (B – D), not to scale (E – G).

FIGURE 12. Morph A of Brachydesmus kalischewskyi Lignau, 1915. A: male (?) from an unspecified locality in Abkhazia, length 15 mm. B: male from Khosta, Krasnodar Prov., Russia. C: male from Pass Mamisoni, W Georgia. Watercolour A after Lignau (1915); pictures B and C by K. Makarov, taken not to scale.

FIGURE 13. Morph A of Brachydesmus kalischewskyi Lignau, 1915: A – C: female from Pass Ertso, South Ossetia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – F: male from Pass Ertso, South Ossetia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. G – H: gonopods of a male from near Batumi, Ajaria, Georgia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal apical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – F by S. Dashdamirov; G – H after Lohmander (1936). Scale bar: 1.0 mm (A – F), not to scale (G – H).

FIGURE 14. Morph B of Brachydesmus kalischewskyi Lignau, 1915: A: male from Yarymdja, E Caucasus Major, Azerbaijan, habitus, lateral view. B: male from Khosrov Nature Reserve, Caucasus Minor, S Armenia, habitus, dorsal view. C: left gonopod of male holotype of B. karawajewi Lohmander, 1928, from Teberda, Karachaevo-Cherkessia, Russia, mesal view. D – E: male from Verkhniy Agdan (now Gandzakar), Idjevan Distr., Armenia (not Georgia, as mistakenly stated by Lohmander 1936), ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Pictures A and B by K. Makarov; C – E after Lohmander (1928, 1936). Not to scale.

FIGURE 15. Morph C of Brachydesmus kalischewskyi Lignau, 1915. A: male from Lower Roki, South Ossetia, habitus, lateral view. B: male from near Krasnaya Polyana, Krasnodar Prov., Russia, habitus, dorsal view. Pictures by K. Makarov, taken not to scale.

FIGURE 16. Morph C of Brachydesmus kalischewskyi Lignau, 1915. A – C: male syntype of B. ferrugineus Lohmander, 1936 from near Borzhomi, Georgia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – F: male syntype of B. ferrugineus Lohmander, 1936 from Novyi Afon, Abkhazia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. G – H: gonopods of syntype of B. ferrugineus from Novyi Afon, Abkhazia, ventral and mesal views, respectively. Designations: en, endomere; ex, exomere; i, mesal apical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – F by S. Dashdamirov; G – H after Lohmander (1936). Scale bar: 1.0 mm (A – F), not to scale (G – H).

FIGURE 18. Morph D of Brachydesmus kalischewskyi Lignau, 1915, male holotype of B. talyschanus Lohmander, 1936: A – C: anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – E: left gonopod, mesal and ventral views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Line drawings A – C by S. Dashdamirov; D – E after Lohmander (1936). Scale bar: 1.0 mm (A – C), not to scale (D – E).

FIGURE 19. Morph E of Brachydesmus kalischewskyi Lignau, 1915, males from near Mestia, Svanetia, Georgia. A – B: habitus, dorsal views. C: leg 7, lateral view. D – E: right gonopod, lateral and mesal views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Pictures A and B by K. Makarov. Scale bars: 0.2 mm (C – E), not to scale (A – B).

FIGURE 21. Morph F of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Agveran, Armenia, anterior part of body, right half of metatergite 10 and caudal part of body, respectively, dorsal views. D – E: male from near Shvanidzor, Megri Mt. Range, Armenia, right gonopod, ventral and mesal views, respectively. F: male from near Badara, Mountainous Karabakh, Azerbaijan, right gonopod, mesal view. Line drawings A – C by S. Dashdamirov. Scale bars: 1.0 mm (A – C), 0.2 mm (D – F).

FIGURE 22. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A: male from Pasture Abago, Caucasian Nature Reserve, Adygea, morph A. B: male from Khosrov Nature Reserve, S Armenia, morph B. C: male from Pass Ertso, South Ossetia, morph A. D: male from Nadirkhanly, Mountainous Karabakh, Azerbaijan, morph F. E: male from Pass Voratan, Zangezur Mt. Range, Armenia, morph B. F: male from near Pass Krestovyi, Georgia, morph B. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bar: 0.2 mm.

FIGURE 23. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Psebe, Krasnodar Prov., Russia, morph A?, lateral, mesal and ventral views, respectively. D – F: male from Zeraboseli, Ajaria, Georgia, morph A?, sublateral, dorsal and ventral views, respectively. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere. Scale bars: 0.1 mm.

FIGURE 24. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Farm Azerbaijan, Talysh Mts, Azerbaijan, morph D, dorsal, ventral and mesal views, respectively. D – F: male from Shishkert, Shikahoh Nature Reserve, Armenia, morph B?, mesal, lateral and ventral views, respectively. Designations: i, mesal, subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bar: 0.1 mm.

FIGURE 25. Examples of gonopods of Brachydesmus kalischewskyi Lignau, 1915. A – C: male from Farm Azerbaijan, Talysh Mts, Azerbaijan, morph D, lateral, mesal and ventral views, respectively. D: male from Arasbaran Wildlife Refuge, East Azerbaijan Prov., Iran, morph D, mesal view. Designations: i, mesal subapical tooth of exomere; e, lateral apical tooth of exomere; l, subapical lobe of exomere. Scale bars: 0.2 mm.

FIGURE 27. Brachydesmus kalischewskyi Lignau, 1915, morph B, males from Pasture Abago and Arkhyz, Adygea, Russia, respectively. A: leg 7, lateral view. B: left gonopod, mesal view. C: leg 7, lateral view. D: left gonopod, mesal view. Scale bars: 1.15 mm (A), 0.2 mm (B and D), 0.1 mm (C).

FIGURE 28. Brachydesmus kvavadzei sp. n. A: male paratype, habitus, dorsal view. B: leg 7, lateral view. C – D: right gonopod, mesal and lateral views, respectively. Designations: en, endomere; ex, exomere; e, apical tooth of exomere. Picture A by K. Makarov, taken not to scale. Scale bars: 0.25 mm (B), 0.1 mm (C – D).

FIGURE 29. Brachydesmus simplex sp. n., male paratype from Sochi, Krasnodar Prov., Russia. A: habitus, dorsal view. B: leg 7, lateral view. C – D: left gonopod, ventral and dorsal views, respectively. E: right gonopod telopodite of male paratype from near Lake Ritsa, Abkhazia, lateral view. F – G: left gonopod of male paratype from Myussera Nature Reserve, Abkhazia, dorsal and ventral views, respectively. Designation: e, lateral apical tooth of exomere. Scale bar: 0.1 mm (B – G), not to scale (A).