Diplomaragnidae (Mikhaljova, 2017)

Mikhaljova, Elena V., 2021, The millipede family Diplomaragnidae Attems, 1907 in the Asian part of Russia with the descriptions of two new genera and seven new species (Diplopoda, Chordeumatida), Zootaxa 5060 (1), pp. 1-44 : 37-39

Key to valid Diplomaragnidae genera and species occurring in the Asian part of Russia

(based only on males)

1(2) Telopodites of anterior gonopods freely positioned regardless of colpocoxites. Mesal sheath processes transformed into a high trapezoidal structure, strongly curved forward ( Fig. 2 )............ Genus Alineuma gen. nov., A. rodionovi sp. nov.

2(1) Telopodites of anterior gonopods positioned inside sheath grooves on posterior surfaces of colpocoxites or in the concave colpocoxites. Mesal sheath processes, if present, different................................................... 3

3(12) Telopodites of anterior gonopods broad or thick all along their length, excluding apical branch, if present............................................................................................... Genus Ancestreuma

4(9) Distal part of anterior gonopod telopodite branched........................................................ 5

5(6) Lateral process of posterior gonopod colpocoxite sheath groove long............................ A. longibrachiatum

6(5) Lateral process of posterior gonopod colpocoxite sheath groove like a plate or absent............................. 7

7(8) Anterior gonopod telopodite with long, subulate branch hidden in the upper part of colpocoxite........... A. subulatum

8(7) Anterior gonopod telopodite with long, tongue-shaped branch not hidden in the upper part of colpocoxite.. A. ramiferum

9(4) Distal part of anterior gonopod telopodite not branched................................................... 10

10(11) Posterior angiocoxal process present............................................................ A. ryvkini

11(10) Posterior angiocoxal process absent........................................................... A. feynmani

12(3) Telopodites of anterior gonopods subflagelliform or flagelliform, or, if broad, then broadened only at base........... 13

13(18) Basal part of anterior gonopod telopodite dilated, distal part acuminate............................. Genus Asiatyla

14(15) Colpocoxite of posterior gonopod with strongly narrowed apex................................. A. communicantis

15(14) Colpocoxite of posterior gonopod with rounded apex..................................................... 16

16(17) Mesal process of the colpocoxite sheath groove with a small apical outgrowth.......................... A. similata

17(16) Mesal process of the colpocoxite without a small apical outgrowth................................... A. sajanica

18(13) Basal part of anterior gonopod telopodite not dilated, distal part flagelliform or subflagelliform.................... 19

19(22) Colpocoxites of posterior gonopods flattened on sides, not sagittally. Lateral sheath process very long and broad ( Fig. 28 ).................................................................................... Genus Orientyla

20(21) Lateral edge of posterior gonopod colpocoxite like a narrow bolster.................................. O. dahurica

21(20) Lateral edge of posterior gonopod colpocoxite like a wide lobe.................................. O. bureyinskaya

22(19) Colpocoxites of posterior gonopods different, if flattened, then frontocaudally. Lateral sheath process not very long and broad........................................................................................... 23

23(32) Posterior gonopod colpocoxite with a large front prominence............................... Genus Sakhalineuma

24(25) Telopodite of anterior gonopod spiralling...................................................... S. basarukini

25(24) Telopodite of anterior gonopod not coiled into a spiral..................................................... 26

26(27) Male coxa 11 with a digitiform process...................................................... S. tuberculatum

27(26) Male coxa 11 without a digitiform process.............................................................. 28

28(29) Lateral process of colpocoxite sheath groove as a large, apically papillate globula..................... S. globuliferum

29(28) Lateral process of colpocoxite sheath groove cylindrical, curved..................................... S. curvatum

30(31) Lateral process of colpocoxite sheath groove broad, with a blunt apex............................... S. molodovae

31(30) Lateral process of colpocoxite sheath groove large, conical, with a pointed apex..................... S. sakhalinicum

32(23) Posterior gonopod colpocoxite without a large front prominence............................................ 33

33(36) Male leg 4 femora with depressed glabrous area, framed by group of setae ( Fig. 18 )................. Litovkia gen. nov.

34(35) Posterior angiocoxal process as a two-lobed lump ( Fig. 21 ). Male body size: 15–16 mm ......... L. anisimovka sp. nov.

35(34) Posterior angiocoxal process laminate, curved in semicircle. Male body size: about 12 mm ........ L. lysaya comb. nov.

36(33) Male leg 4 femora without depressed glabrous area, framed by group of setae.................................. 37

37(68) Gonopod anterior angiocoxal process long, subflagelliform to flagelliform.................................... 38

38(59) Gonopod anterior angiocoxal processes passing through a foramen, between colpocoxites along midline or each process placed inside a fold of colpocoxite ( Fig. 5 )............................................... Genus Diplomaragna

39(44) Male coxa 11 ventrofrontally with papillate field and rounded outgrowth distally................................ 40

40(41) Posterior angiocoxal processes with apical and subapical strong teeth and subbasal papillate rounded outgrowth, medially contiguous.............................................................................. D. konyukhovi

41(40) Posterior angiocoxal processes different: elongted vertically, at base with mesal outgrowths meeting in midline ( Fig. 13 ).......................................................................................... D. provecta

42(43) Edges of front colpocoxital folds set horizontally to body axis, posterior angiocoxal process with a pointed apex.............................................................................................. D. kedrovaya

43(42) Edges of front colpocoxital folds set obliquely to body axis, posterior angiocoxal process with a blunt apex ( Fig. 15 )........................................................................................... D. yakovlevka

44(39) Male coxa 11 ventrofrontally without papillate field and rounded outgrowth distally............................. 45

45(46) Apex of gonopod posterior angiocoxal process expanded, with large and small teeth meeting in midline..... D. terricolor

46(45) Apex of gonopod posterior angiocoxal process different....................................................47

47(48) Posterior gonopod colpocoxite with a broad blunt tip................................................ D. ganini

48(47) Posterior gonopod colpocoxite with a narrow, often pointed, sometimes unciform tip............................ 49

49(54) Posterior angiocoxal process like a caudally convex plate...................................................50

50(51) Posterior angiocoxal process with a small process-tooth subbasally ( Fig. 9 )....................... D. dentifer sp. nov.

51(50) Posterior angiocoxal process without a small process-tooth subbasally........................................ 52

52(53) Distal part of the colpocoxite flagelliform, tip hooked........................................... D. zimoveinaya

53(52) Distal part of the colpocoxite not flagelliform, slightly curved caudally, tip hooked ( Fig. 3 ).......... D. budilovi sp. nov.

54(49) Posterior angiocoxal process not like a caudally convex plate............................................... 55

55(56) Colpocoxites of posterior gonopods strongly curved caudad and deeply emarginate at each mesal edge, forming a large cordiform foramen........................................................................ D. anuchino

56(55) Colpocoxites of posterior gonopods either not curved strongly caudad or only their tips curved; no deep emargination at each mesal colpocoxital edge........................................................................ 57

57(58) Posterior angiocoxal process curved forward distally as well as with a long, slender, median spine....... D. dalnegorica

58(57) Posterior angiocoxal process not curved forward distally as well as without a long, slender median spine............ 59

59(38) Gonopod anterior angiocoxal processes either piercing the colpocoxites or sheathed on anterior face of colpocoxites.................................................................................... Genus Maritimosoma

60(61) Anterior angiocoxal process positioned on stick-shaped outgrowth of angiocoxite, as result it looks divided into two unequal parts........................................................................................ M. antis

61(60) Anterior angiocoxal process not positioned on stick-shaped outgrowth of angiocoxite, as result it not looks divided into two unequal parts..................................................................................... 62

62(63) Anterior angiocoxal process piercing the colpocoxite............................................... M. turova

63(62) Anterior angiocoxal process sheathed on anterior face of colpocoxite..........................................64

64(65) Posterior angiocoxal process not like a caudad convex plate........................................ M. reductum

65(64) Posterior angiocoxal process like a caudad convex plate................................................... 66

66(67) Posterior angiocoxal process with a long, slender, distal process....................................... M. piceum

67(66) Posterior angiocoxal process with an apical and a median pointed horn strongly separated.............. M. schawalleri

68(37) Gonopod anterior angiocoxal process shorter, of a different shape............................................69

69(82) Colpocoxite sheath groove without evident processes........................................Genus Pacifiosoma

70(79) Colpocoxites completely or only posteriorly densely covered with papillae and/or pointed spinules................. 71

71(76) Posterior angiocoxal process without clawlike outgrowth but with rounded pappilate/spinulate outgrowth............ 72

72(73) Colpocoxite only posteriorly densely covered with papillae.................................. P. shabalini sp. nov.

73(72) Colpocoxite completely densely covered with pointed spinules and papillae................................... 74

74(75) Apex of colpocoxite pointed................................................................... P. koiensis

75(74) Apex of colpocoxite not pointed............................................................... P. asperum

76(71) Posterior angiocoxal process with clawlike outgrowth..................................................... 77

77(78) Mesal margin of gonopod colpocoxite drawn into a triangular plate ( Fig. 56 )................. P. triangulatum sp. nov.

78(77) Mesal margin of gonopod colpocoxite not drawn into a triangular plate ( Fig. 32 ).................. P. acutum sp. nov.

79(70) Colpocoxites completely or only posteriorly not covered with papillae and/or pointed spinules.................... 80

80(81) Anterior angiocoxal processes of posterior gonopods present ( Fig. 45 )................................. P. cristofer

81(80) Anterior angiocoxal processes of posterior gonopods absent.......................................... P. kuruma

82(69) Colpocoxite sheath groove with evident (at least with mesal sheath process) processes........................... 83

83(94) Posterior angiocoxal process of posterior gonopod medium-sized, very small or absent. Colpocoxite (sub)cyathiform in shape................................................................................. Genus Shearia

84(89) Mesal sheath processes fused into a large cup-shaped structure.............................................. 85

85(86) Posterior angiocoxal process of posterior gonopod rather well-developed............................... S. oiskaya

86(85) Posterior angiocoxal process of posterior gonopod small, almost absent........................................87

87(88) Anterior angiocoxal process very large, broad, with a small lateral projection....................... S. shushenskaya

88(87) Anterior angiocoxal process different: bifurcated................................................ S. khakassica

89(84) Mesal sheath processes fused into a small or middle size cup-shaped structure.................................. 90

90(91) Apex of the colpocoxite with a subquadrate excavation. Telopodites of anterior gonopods very long......... S. rybalovi

91(90) Apex of the colpocoxite different. Telopodites of anterior gonopods shorter.....................................92

92(93) Anterior angiocoxal process very large, broad, with a lateral dentiform projection....................... S. teletskaya

93(92) Anterior angiocoxal process small, different............................................................ 94

94(95) Posterior gonopod angiocoxite in anterior view with one outgrowth................................... S. calycina

95(94) Posterior gonopod angiocoxite in anterior view with two outgrowths................................. S. densecava

96(83) Posterior angiocoxal process of posterior gonopod large. Colpocoxite different.................... Genus Altajosoma

97 (98) Basal half of colpocoxite in anterior view with longitudinal crest................................... A. kultukensis

98(97) Basal half of colpocoxite in anterior view without longitudinal crest......................................... 99

99(102) Lateral process of colpocoxite sheath groove very long, positioned basally................................... 100

100(101) Posterior angiocoxal process with a large lateral and three smaller mesal teeth........................ A. deplanatum

101(100) Posterior angiocoxal process different: large, curved forward, with pointed apex....................... A. corniferum

102(99) Lateral process of colpocoxite sheath groove not very long, not basally positioned............................. 103

103(104) Male coxae 12 enlarged, trapezoidal, with small cones on posterior faces............................... A. coxale

104(103) Male coxae 12 different............................................................................ 105

105(106) Apex of the posterior angiocoxal process without teeth............................................ A. shilenkovi

106(105) Apex of the posterior angiocoxal process with teeth..................................................... 107

107(108) Lateral process of colpocoxite sheath groove as large blade with finger-shape ventrally extending outgrowth. A. baltyrgan

108(107) Lateral process of colpocoxite sheath groove different................................................... 109

109(110) Lateral process of colpocoxite sheath groove strongly projecting beyond colpocoxite, well visible in anterior view, its tip swollen................................................................................ A. golovatchi

110(109) Lateral process of colpocoxite sheath groove not protruding considerably beyond colpocoxite, almost invisible in anterior view, its apex different............................................................................ 111

111(116) Lateral process of colpocoxite sheath groove with a laterally cupped lamella.................................. 112

112(113) None of the five teeth on posterior angiocoxal process set well apart from the others.................... A. kemerovo

113(112) One of the five teeth on posterior angiocoxal process set well apart from the others............................ 114

114(115) Anterior angiocoxal process of posterior gonopods short.......................................... A. bakurovi

115(114) Anterior angiocoxal process of posterior gonopods long...................................... A. longiprocessum

116(111) Lateral process of colpocoxite sheath groove without a laterally cupped lamella........................ A. katunicum

Figures

FIGURES 1–2. Alineuma rodionovi sp. nov., male holotype. 1, gonopods, front view; 2, gonopods, caudal view, right femur of posterior gonopod telopodite is twisted; ms, trapezoidal structure; b, hyaline fimbriate branch; ap, anterior angiocoxal process; pp, posterior angiocoxal process; at, anterior gonopod telopodite; sg, subglobule of posterior angiocoxite. Scale in mm.

FIGURES 28–31. Orientyla dahurica (Gerstfeldt, 1859), male. 28, gonopods, caudal view; 29, gonopods, front view; 30, distal parts of colpocoxites, front view; 31, apex of colpocoxite, front view; lp, lateral process of sheath groove; at, anterior gonopod telopodite; e, edge of sheath groove; pp, posterior angicoxal process; cx, coxosternum. Scales: 20 μm (Fig. 31), 100 μm (Fig. 29), 200 μm (Fig. 28), 0.5 mm (Fig. 30).

FIGURES 17–22. Litovkia anisimovka sp. nov., male paratype. 17, distal part of leg 4; 18, leg 4 without coxa and trochanter; 19, leg pair 11, caudal view; 20, coxae and trochanters 11, caudal view; 21, gonopods, caudal view, telopodites of posterior gonopods broken; 22, distal part of gonopods, caudal view; r, ridges-filaments; pp, posterior angicoxal process; ap, bifid anterior angiocoxal process; at, anterior gonopod telopodite; a, glabrous area. Scales: 20 μm (Figs 17, 22), 100 μm (Figs 18–21).

FIGURES 3–7. Diplomaragna budilovi sp. nov., male holotype. 3, 7, gonopods, caudal view; 4, posterior angiocoxal processes and colpocoxites, caudal view; 5, gonopods, front view; 6, basal part of gonopods, caudal view; c, horn; pp, posterior angicoxal process; ap, anterior angiocoxal process; cx, coxosternum; at, anterior gonopod telopodite. Scales: 20 μm (Fig. 6), 100 μm (Figs 3–5), 0.5 mm (Fig. 7).

FIGURES 13–14. Diplomaragna provecta Mikhaljova, 2005, male. 13, gonopods, caudal view; 14, central part of colpocoxite; pp, posterior angicoxal process; pt, posterior gonopod telopodite; cl, colpocoxites. Scales: 20 μm (Fig. 14), 100 μm (Fig. 13).

FIGURES 15–16. Diplomaragna yakovlevka Shear, 1990, male. 15, gonopods, caudal view; 16, posterior angiocoxal processes, caudal view; sg, subglobule of posterior angiocoxite. Scales: 20 μm (Fig. 16), 100 μm (Fig. 15).

FIGURES 8–12. Diplomaragna dentifer sp. nov., male, female paratypes. 8, gonopods, caudal view; 9, 10, posterior angiocoxal processes; 11, gonopods, front view; 12, vulvae, ventral view; pp, posterior angicoxal process; t, process-tooth; ap, anterior angiocoxal process. Scales: 20 μm (Figs 9–10), 100 μm (Figs 8, 11–12).

FIGURES 56–57. Pacifiosoma triangulatum sp. nov., male holotype. 56, gonopods, caudal view; 57, gonopods, front view; pp, posterior angicoxal process; c, clawlike outgrowth; n, triangular plate; ap, anterior angiocoxal process; cx, coxosternum. Scale in mm.

FIGURES 32–36. Pacifiosoma acutum sp. nov., male holotype. 32, 36, gonopods, caudal view, right colpocoxite broken; 33, posterior angiocoxal processes; 34, distal parts of anterior gonopod telopodite and colpocoxite; 35, external edge of colpocoxite; pp, posterior angicoxal process; n, external edge of colpocoxite; at, anterior gonopod telopodite; c, clawlike outgrowth. Scales: 10 μm (Figs 34–35), 20 μm (Fig. 33), 100 μm (Fig. 32), 0.5 mm (Fig. 36).

FIGURES 45–48. Pacifiosoma cristofer (Mikhaljova, 1993), males from Gassi Lake basin (Figs 45–46) and Bolshekhekhtsirsky Nature Reserve (Figs 47–48). 45, 47, gonopods, front view; 46, 48, distal parts of colpocoxites; ap, anterior angiocoxal process; cl, colpocoxites. Scales: 20 μm (Figs 46, 48), 100 μm (Figs 45, 47).