Plusioglyphiulus, Silvestri, 1923

Golovatch, Sergei I., Geoffroy, Jean-Jacques, Mauriès, Jean-Paul & Vandenspiegel, Didier, 2011, The millipede genus Plusioglyphiulus Silvestri, 1923 in Thailand (Diplopoda, Spirostreptida, Cambalopsidae) 2940, Zootaxa 2940 (1), pp. 1-63 : 60-61

Key to species of Plusioglyphiulus :

1. Crests on collum not differentiated, arranged in two transverse rows ( Figs 5A, D ), just like in typical Glyphiulus species. Gonopods also much like in typical Glyphiulus species ( Figs 7F, G ), but each posterior gonopod coxite with a distinct apical ampulla devoid of a nearby flagellum ( Fig. 7G )..................................................... P. antiquior

– Crests on collum markedly differentiated, usually broken and arranged in three transverse rows ( Figs 1A, D , 8A, D ). Anterior gonopods usually not shield-shaped like in typical Glyphiulus species , but more complex and typically divided into a pair of anterior and posterior coxosternal processes ( Figs 11G , 15C , 18C, D ). Each posterior gonopod coxite with a distinct apical ampulla either supplied with a nearby flagellum or not; when supplied, then flagellum nearly always simple ( Figs 11H , 18E, F ), very rarely plumose or serrate ( Fig. 15D )................................................................ 2

2. At least some crests on collum complete ( Figs 8A, D , 28A, D , 39A, D , 51A, D ).................................... 3

– No complete crests on collum, all interrupted ( Figs 1A, D , 12A, D , 16A, D )....................................... 9

3. Collum with 3(4)+3(4) complete crests..................................................................... 4

– Collum with 5(6)+5(6) or 1+1 complete crests............................................................... 6

4. Complete crests on collum: 2, 3 and 4, paramedians incomplete. Sabah, Malaysia........................ P.macfarlanei

– Complete crests on collum always including paramedians...................................................... 5

5. Collum with 3+3 complete crests ( Figs 28A, D ). Gonopods as in Figs 30B–D , 31C–E . North-central Thailand....... P.tham

– Collum with 4+4 complete crests ( Figs 51A, D ). Gonopods as in Figs 52F, G . Southern Vietnam.............. P. ampullifer

6. Only paramedians on collum complete. Carinotaxy formula of midbody segments, 3/3+I/i+3/3/3+m/m/m. Thailand........ 7

– Collum with 5(6)+5(6) complete crests. Carinotaxy formula of midbody metaterga different. Borneo................... 8

7. Gonopods as in Figs 41G, H.................................................................... P . puttakun

– Gonopods as in Figs 11G, H.................................................................... P . sutchariti

8. Carinotaxy formula of midbody segments, 3/3+I/i+2/2+m/m. Cave in Sarawak, Malaysia................ P. cavernicolus

– Carinotaxy formula of midbody segments, 4/4+I/i+3/3/3+m/m. Cave in Kalimantan, Indonesia................ P. bedosae

9. Carinotaxy pattern of collum isostichic (i.e. devoid of intercalary tubercles or crests). Laos.................... P.steineri

– Carinotaxy pattern of collum mixostichic.................................................................. 10

10. Carinotaxy formula of midbody metaterga 3/3+I/i+3/3+m/m. Anterior gonopods with a prominent, spiniform, median, sternal process. Caves in Malaya.................................................................... P. grandicollis

– Carinotaxy formulae of midbody metaterga different. Anterior gonopods without sternal process...................... 11

11. Carinotaxy formula of midbody metaterga, 2/2+I/i+1/1+m/m. Male leg 1 with a relatively short central hook ( Fig. 4C ). Male leg 2 moderately enlarged ( Fig. 4D ). Anterior gonopods shield-shaped ( Figs 3D , 4G ), like in Glyphiulus species. Posterior gonopods much like in P. antiquior , but supplied with evident, bare flagella on top ( Figs 3B, C , 4H )............. P. panhai

– Carinotaxy formulae of midbody metaterga different. Male leg 1 with a strong, long, central hook ( Fig. 11C ). Male leg 2 moderately enlarged ( Fig. 11D ). Anterior gonopods much more elaborate, with two pairs of prominent processes ( Figs 11G , 15C , 18C ). Posterior gonopods different, telopodites rudimentary to missing......................................... 12

12. Second transverse row of tubercles on midbody segments intercalary, apparently developed only towards onset of sexual maturity, formula 3/3+I/i+3/4/3+m//m. Epiproct short. Legs more or less as long as body diameter. Borneo................. 13

– Dorsal tubercles or crests arranged in clear-cut longitudinal rows, pattern isostichic ( Figs 12B, E , 16B, E , 20B, E ). Epiproct longer ( Figs 12C, F , 13D, E , 16C, F , 20C, F ). Legs usually closer to being half as long as body diameter ( Figs 13F , 17E , 21F ). ................................................................................................... 15

13. Body length 43–50 mm, width 2.0– 2.1 mm; segments 64–70p+2–1a+ T. Collum with a distinct transverse impression near anterior 1/3. Carinotaxy formula of midbody segments 2/2+I/i+3/4/3+m//m. Paraprocts flattened. Kalimantan .... P. pallidior

– Body length up to about 30 mm, width 1.0– 1.6 mm; segments up to 53p+1a+T. Collum not particularly impressed near anterior 1/3. Carinotaxy formula of midbody segments 3/3+I/i+3/4(5)/3+m//m. Paraprocts regularly convex................... 14

14. Brown spots present dorsad of ocellaria. Epiproct very short, roundly subtrapeziform. Anterior gonopods without cxp2. Lateral arms of posterior gonopods rudimentary. Sarawak.................................................... P. hoffmani

– Brown spots absent dorsad of ocellaria. Epiproct a little longer, regularly rounded. Anterior gonopods swollen, stout, with both cxp1 and cxp2 present. Lateral arms of posterior gonopods long and spiniform. Kalimantan ..................... P. similis

15. General coloration very dark brown to blackish (fading to reddish after long conservation in alcohol). Paraprocts with a distinct, median, ridge-like elevation. Cambodia........................................................ P. dubius

– General coloration lighter, usually yellow-brown to brown. Paraprocts flat medially................................ 16

16. Paraprocts strongly flattened. Each posterior gonopod with a long lateral spine.......................... P. deharvengi

– Paraprocts regularly convex ( Figs 12C , 16C , 20C ). Posterior gonopods without long spine laterally.................... 17

17. Carinotaxy of midbody metaterga, 3/3+I/i+3/3/3+m/m/m, changing 4/4+I/i+3/3/3+m/m toward caudal third of body...... 18

– Basic carinotaxy of midbody metaterga, 3/3+I/i+3/3/3+m/m/m, remaining unchanged toward telson.................. 23

18. Anterior gonopods as in Figs 14D, E , 15C ; posterior gonopods with plumose flagella on top ( Fig. 15D ).......................................................................................................... P. likhitrakarni

– Anterior gonopods different; flagella on posterior gonopods either simple or absent................................ 19

19. Anterior gonopods with cxp1 considerably longer than cxp2 ( Figs 22C–E , 23G ); flagella on to posterior gonopods very short, telopodites missing, but coxite with a conspicuous frontomedial arm ( Fig. 23H )............................. P. erawan

– Anterior gonopods with cxp1 usually shorter than to about as long as cxp2; posterior gonopods different, a frontomedian arm less conspicuous or missing............................................................................ 20

20. Anterior gonopods as in Figs 26D, E , 27C ; posterior gonopods shield-like, front arms short ( Figs 26F , 27D ).......... P.wat

– Both gonopod pairs different............................................................................ 21

21. Anterior gonopods as in Figs 18C, D , 19F ; posterior gonopods with very long flagella on top ( Figs 18E, F , 19G ).................................................................................................... P.pimvichaiae

– Both gonopod pairs different............................................................................ 22

22. Anterior gonopods with rounded tips of cxp1 ( Figs 34A–D , 35F ); posterior gonopods virtually devoid of flagella ( Figs 34E, F , 35G )........................................................................................... P. phra

– Anterior gonopods with acuminate cxp1 ( Fig. 50F ); posterior gonopods more elaborate, flagella present ( Fig. 50G )...................................................................................................... P. bessoni

23. Brown spots present dorsad of ocellaria. Anterior gonopods with very short and lobe-like cxp1. Posterior gonopods without arms. Cambodia................................................................................ P. boutini

– Brown spots absent dorsad of ocellaria. Anterior gonopods with both cxp1 and cxp2 equally well developed. Posterior gonopods with or without arms............................................................................... 24

24. Anterior gonopods as in Figs 47G, H ; posterior gonopods more elaborate, with clear telopodite remains ( Fig. 47I )..................................................................................................... P. samakkee

– Both gonopod pairs different............................................................................ 25

25. Anterior gonopods with both cxp1 and cxp2 bifid; posterior gonopods with rudimentary arms, devoid of flagella. Laos.................................................................................................. P. foveatus

– Anterior gonopods with cxp1 uncinate while cxp2 tip either digitiform or expanded; posterior gonopods without arms, but with flagella. Thailand..................................................................................... 26

26. Anterior gonopods with digitiform cxp2 ( Fig. 38E ); posterior gonopods as in Figs 38F, G...................... P . jaydee

– Anterior gonopods with subtriangular tips of cxp2 ( Fig. 44G ); posterior gonopods as in Fig. 44H................ P .saksit

Figures

FIGURE 1. Plusioglyphiulus panhai sp. n., male paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A, C–F, 0.2 mm; B, 0.5 mm.

FIGURE 3. Plusioglyphiulus panhai sp. n., male paratype. A: enlarged claw with a basal setoid filament, subdorsal view. B: both gonopod pairs in situ, caudal view. C: enlarged right half of same, caudal view. D: anterior gonopods in situ, caudal view. Scale bars: A, 0.01 mm; B and D, 0.1 mm; C, 0.05 mm.

FIGURE 4. Plusioglyphiulus panhai sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, caudal view. D: leg 2, caudal view. E: leg 3, caudal view. F: midbody leg. G: anterior gonopods, caudal view. H: posterior gonopods, caudal view. te: telopodite; cxp: coxosternal process; lo: ventrolateral lobe on coxosternum. Scale bars: A–F, 0.4 mm, G and H, 0.2 mm.

FIGURE 5. Plusioglyphiulus antiquior sp. n., female paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–F, 0.2 mm.

FIGURE 7. Plusioglyphiulus antiquior sp. n., male holotype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, subcaudal view. D: leg 2, caudal view. E: midbody leg. F: anterior gonopods, caudal view. H: posterior gonopods, front view. te: telopodite; cxp: coxosternal process. Scale bar: 0.2 mm.

FIGURE 8. Plusioglyphiulus sutchariti sp. n., female paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–F, 0.2 mm.

FIGURE 11. Plusioglyphiulus sutchariti sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, front view. D: leg 2, caudal view. E: leg 3, caudal view. F: midbody leg. G: anterior gonopods, front view. H: right posterior gonopod, caudal view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A– F, 0.4 mm; G and H, 0.2 mm.

FIGURE 12. Plusioglyphiulus likhitrakarni sp. n., male paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–F, 0.2 mm.

FIGURE 13. Plusioglyphiulus likhitrakarni sp. n., male paratype. A: anterior part of body, ventral view. B: gnathochilarium, ventral view. C: midbody segments, ventral view. D: posterior part of body, ventrocaudal view. E: same, dorsocaudal view. F: cross-section of a midbody segment, caudal view. Scale bars: A, C–F, 0.2 mm; B, 0.1 mm.

FIGURE 14. Plusioglyphiulus likhitrakarni sp. n., male paratype. A: segment 7, sublateral view. B: enlarged ozopore region, lateral view. C: enlarged claw, sublateral view; D: anterior gonopods, front view. E: same, ventrocaudal view. F: posterior gonopods, caudolateral view. Scale bars: A, B, D and E, 0.1 mm; C, 0.01 mm; F, 0.05 mm.

FIGURE 15. Plusioglyphiulus likhitrakarni sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: left anterior gonopod, front view. D: posterior gonopods, front view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: 0.2 mm.

FIGURE 16. Plusioglyphiulus pimvichaiae sp. n., male paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–C, E and F, 0.2 mm; D, 0.5 mm.

FIGURE 17. Plusioglyphiulus pimvichaiae sp. n., male paratype. A: anterior part of body, frontoventral view. B: gnathochilarium, ventral view. C: midbody segments, ventral view. D: posterior part of body, ventral view. E: cross-section of a midbody segment, caudal view; F: enlarged ozopore region, lateral view. Scale bars: A, 0.5 mm; B–E, 0.2 mm; F, 0.1 mm.

FIGURE 18. Plusioglyphiulus pimvichaiae sp. n., male paratype. A: legs 1–3, caudal view. B: enlarged claw, dorsolateral view. C: anterior gonopods, front view. D: same, caudal view. E: posterior gonopods, caudal view. F: same, ventrocaudal view. Scale bars: A, 0.1 mm; B, 0.01 mm; C−F, 0.05 mm.

FIGURE 19. Plusioglyphiulus pimvichaiae sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, subfront view. D: leg 2, caudal view. E: leg 3, front view. F: anterior gonopods, caudolateral view. G: posterior gonopods, subcaudal view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–E, 0.4 mm; F and G, 0.2 mm.

FIGURE 20. Plusioglyphiulus erawan sp. n., male paratype from Cave Erawan. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–E, 0.5 mm; F, 0.2 mm.

FIGURE 21. Plusioglyphiulus erawan sp. n., male paratypes from Cave Erawan. A: anterior part of body, ventrolateral view. B: gnathochilarium, ventral view. C: midbody segments, ventral view. D: posterior part of body, ventral view. E: same, caudal view. F: cross-section of a midbody segment, caudal view; G: enlarged ozopore region, lateral view. Scale bars: A and F, 0.5 mm; B–E, 0.2 mm; G, 0.01 mm.

FIGURE 22. Plusioglyphiulus erawan sp. n., male paratypes from Cave Erawan. A: enlarged claw, lateral view. B: both gonopod pairs, front view. C: same, caudal view. D: same enlarged, caudoventral view. E: same, ventrolateral view. Scale bars: A, 0.02 mm; B−E, 0.1 mm.

FIGURE 23. Plusioglyphiulus erawan sp. n., male paratype from Cave Erawan. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, front view. D: leg 2, front view. E: leg 3, front view. F: midbody leg, lateral view. G: anterior gonopods, caudal view. H: posterior gonopods, front view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: 0.2 mm.

FIGURE 26. Plusioglyphiulus wat sp. n., male paratype. A: gnathochilarium, ventral view. B: enlarged claw, lateral view. C: legs 1−3, subventral view. D: anterior gonopods, caudal view. E: same, front view. F: posterior gonopods, caudal view. Scale bars: A, 0.2 mm; B, 0.02 mm; C−E, 0.1 mm; F, 0.05 mm.

FIGURE 27. Plusioglyphiulus wat sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: left anterior gonopod, caudal view. D: posterior gonopods, front view. Scale bar: 0.2 mm.

FIGURE 28. Plusioglyphiulus tham sp. n., male paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A and B, 0.5 mm; C–F, 0.2 mm.

FIGURE 30. Plusioglyphiulus tham sp. n., male paratype. A: enlarged claw, ventrolateral view. B: anterior gonopods, frontoventral view. C: same, caudal view. D: same, caudoventral view. Scale bars: A, 0.02 mm; B−D, 0.1 mm.

FIGURE 31. Plusioglyphiulus tham sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: right anterior gonopod, caudal view. D: left posterior gonopod, caudal view. E: same, front view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A and B, 0.4 mm; C and D, 0.2 mm.

FIGURE 34. Plusioglyphiulus phra sp. n., male paratype. A: anterior gonopods, front view. B: same, ventral view. C: same, ventrolateral view. D: same, ventrocaudal view. E: posterior gonopods, ventrofront view. F: same, front view. Scale bars: A, C and F, 0.1 mm; B, D and E, 0.05 mm.

FIGURE 35. Plusioglyphiulus phra sp. n., male paratype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, caudal view. D: leg 2, caudal view. E: midbody leg, lateral view. F: anterior gonopods, frontolateral view. G: posterior gonopods, caudolateral view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–E, 0.4 mm; F and G, 0.2 mm.

FIGURE 38. Plusioglyphiulus jaydee sp. n., male paratype. A: gnathochilarium, ventral view. B: leg 1, caudal view. C: leg 2, caudal view. D: leg 3, front view. E: anterior gonopods, front view. F: posterior gonopods, front view. G: same, caudal view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–D, 0.4 mm; E–G, 0.2 mm.

FIGURE 39. Plusioglyphiulus puttakun sp. n., female paratype. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. Scale bars: A–F, 0.2 mm.

FIGURE 41. Plusioglyphiulus puttakun sp. n., holotype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, sublateral view. D: leg 2, front view. E: leg 3, caudal view. F: midbody leg, lateral view. G: anterior gonopods, caudal view. H: posterior gonopods, front view. I: same, lateral view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–F, 0.4 mm; G–I, 0.2 mm.

FIGURE 44. Plusioglyphiulus saksit sp. n., holotype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, lateral view. D: leg 2, caudal view. E: leg 3, caudal view. F: midbody leg, lateral view. G: anterior gonopods, front view. H: posterior gonopods, caudal view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–F, 0.4 mm; G and H, 0.2 mm.

FIGURE 47. Plusioglyphiulus samakkee sp. n., holotype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 1, sublateral view. D: leg 2, caudal view. E: leg 3, caudal view. F: midbody leg, lateral view. G: anterior gonopods, caudolateral view. H: right anterior gonopod, mesal view. I: posterior gonopods, front view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–F, 0.4 mm; G–I, 0.2 mm.

FIGURE 50. Plusioglyphiulus bessoni Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009, male topotype. A: antenna, lateral view. B: gnathochilarium, ventral view. C: leg 2, caudal view. D: leg 3, caudal view. E: midbody leg, lateral view. F: anterior gonopods, front view. G: posterior gonopods, caudal view. te: telopodite; cxp1: anterior coxosternal process; cxp2: posterior coxosternal process. Scale bar: A–E, 0.4 mm; F and G, 0.2 mm.

FIGURE 51. Plusioglyphiulus ampullifer Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009, male from Cat Tien. A: anterior part of body, lateral view. B: midbody segments, lateral view. C: posterior part of body, lateral view. D: anterior part of body, dorsal view. E: midbody segments, dorsal view. F: posterior part of body, dorsal view. G: anterior part of body, ventral view. H: posterior part of body, ventral view. I: gnathochilarium, ventral view. J: cross-section of a midbody segment, caudal view. K: enlarged ozopore region, lateral view. L: leg 1, lateral view. M: leg 2, caudal view. Scale bars: A–E, G and J, 0.5 mm; F, H, I and M, 0.2 mm; K and L, 0.1 mm.

FIGURE 52. Plusioglyphiulus ampullifer Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009, male from Cat Tien. A: segment 3 with legs 3, front view. B: midbody leg, lateral view. C: enlarged claw, lateral view. D: ventral parts of segments 6 and 7, ventrolateral view. E: cross-section of segment 7, caudal view. F: both gonopod pairs, ventral view. G: same, caudal view. H: same, front view. Scale bars: A and E, 0.5 mm; B, D and F–H, 0.1 mm; C, 0.02 mm.