Haplodesmidae, Cook, 1895

Golovatch, Sergei, Geoffroy, Jean-Jacques, Mauriès, Jean-Paul & VandenSpiegel, Didier, 2009, Review of the millipede family Haplodesmidae Cook, 1895, with descriptions of some new or poorly-known species (Diplopoda, Polydesmida), ZooKeys 7 (7), pp. 1-53 : 46-49

A key to recognizable genera and species of Haplodesmidae :

1 Body with 20 segments, pyrgodesmid-like (collum flabellate, covering the head from above; metaterga with three transverse rows of tubercles; paraterga well-developed, strongly declivous, porosteles often present), incapable of volvation. Gonopods rather simple, unciform; coxae with only a few ventrolateral setae; telopodites basically uniramous, only sometimes with an inconspicuous, lateral outgrowth at midway, always devoid of a separate solenomere branch; seminal groove terminating distally or apically, hairpad often present ( Prosopodesmus )............................................................................... 2

– Body with 18, 19 or 20 segments, vermiform (without clear paraterga on most body segments, i.e. incapable of volvation) or “doratodesmid” (with very evident paraterga, mostly capable of tight volvation). Gonopods either simplified (= regressed) or more elaborate, often bi- or triramous................ 4

2 Gonopod telopodite with a seminal groove terminating with a hairpad distally (i.e. acropodite rather large, slightly dentate)....................................... 3

– Gonopod telopodite with a seminal groove terminating apically, devoid of hairpad (i.e. acropodite virtually absent) ................ Prosopodesmus sinuatus

3 Gonopod telopodite with a distinct lateral outgrowth at about midway; ozopores absent from paraterga 6 .............................. Prosopodesmus jacobsoni

– Gonopod telopodite devoid of a lateral outgrowth at midway; ozopores present on porosteles on paraterga 6 .................... Prosopodesmus panporus

4 Body subcylindrical, not capable of volvation, with paraterga 2 rather well developed, but subsequent ones represented mostly by lateral swellings. Gonopods simplified ....................................................................................... 5

– Body “doratodesmid”, mostly capable of complete conglobation, with paraterga 2 always very strongly enlarged laterally, all following paraterga more or less strongly declivous while collum somewhat reduced. Gonopods usually rather elaborate......................................................................................... 10

5 Body with 19 (♂) or 20 (♀) segments, collum and all following metaterga with abundant setation in part represented by long, bisegmented, tactile setae. Gonopods especially simple; telopodite = solenomere sometimes with a lateral outgrowth at midway; seminal groove terminating subapically and devoid of hairpad ( Cylindrodesmus ) ............................................................. 6

– Body with 19 or 20 segments regardless of sex, collum and all following metaterga without abundant setation, usually tuberculate. Gonopods aberrant ........... 7

6 Gonopod telopodite virtually uniramous .............. Cylindrodesmus hirsutus

– Gonopod telopodite evidently biramous in distal half.................................. .............................................................................. Cylindrodesmus villosus

7 Body with 19 segments, paraterga 2 rather well developed, but subsequent ones mostly represented by lateral swellings. Collum still large, covering the head from above, with 4-5 transverse rows of setigerous tubercles or pits. Following paraterga with three rows of similar tubercles or pits. Gonopods with poorly setose gonocoxae and a considerably shortened prefemoral+femoral (= setose) part, deeply biramous thereafter, with a long and flagelliform solenomere devoid of hairpad ( Helodesmus ) .......................................................... 8

– Body with 20 segments, paraterga 2 rather well developed, subsequent ones represented by lateral swellings at most; tergal trichome wanting. Collum small. Gonopod coxae virtually bare and reduced; telopodites strongly geniculate and with no traces of either a coxal cannula or a seminal groove ( Atopogonus )..................................................................................................... 9

8 Metaterga behind collum with three transverse rows of short setae borne on small tubercles. Gonopod acropodite a long and simple branch. Java ............ ..................................................................................... Helodesmus porosus

– Metaterga behind collum with three transverse rows of short setae in microsetose pits. Gonopod acropodite a long and bifid branch. Sumatra ............... ................................................................................... Helodesmus parvulus

9 Gonopod acropodite deeply and evidently biramous, apex finely fringed in part............................................................................. Atopogonus baccatus

– Gonopod acropodite compact, vaguely biramous, apex not fringed .............. ................................................................................ Atopogonus bucculatus

10 Gonopod telopodite usually stout and strongly enlarged laterally towards end of femorite ( Figs 29B, C ), with or without a short solenomere branch thereafter; acropodite varying from absent to well-developed ( Doratodesmus ).... 11

– Gonopod telopodite usually slender, not enlarged towards the end of femorite, but with a more or less distinct process or outgrowth laterally ( Figs 4B, C ), opposite recurvature point of seminal groove; solenomere thereafter taking up most of telopodite, sometimes elaborate; seminal groove terminating distally to subapically, with or without a hairpad; acropodite small to nearly absent ( Eutrichodesmus )............................................................................. 16

11 Some metaterga with very evident mid-dorsal processes or outgrowths ..... 12

– Metaterga without mid-dorsal outgrowths ................................................ 13

12 Metaterga 4-19 with a mid-dorsal subtriangular process, gradually increasing in size towards (to become largest on) metatergum 18. Java .......................... ............................................................................... Doratodesmus armatus

– Metaterga before 15 th and after 18 th (penultimate) without mid-dorsal outgrowths; metaterga 15 and 16 each with a small subtriangular process, metatergum 17 with a particularly prominent, rounded, central process. Sumatra ....................................................................... Doratodesmus analdes

13 Most metaterga with 2-3 transverse rows of flat bosses. Gonopod telopodite very simple, femorite very short and stout (much like in Helodesmus ). New Guinea...................................................................................................... 14

– Most metaterga with three transverse rows of distinct conical tubercles. Gonopod telopodite more elaborate ( Figs 29B, C )......................................... 15

14 Male with 18, female with 19 body segments. Gonopod femorite very stout, divided apically into a short, subunciform solenomere and a simple, knifeshaped acropodite .................................................... Doratodesmus pholeter

– Both sexes with 18 body segments. Gonopod femorite turning apically into a rather short, subunciform solenomere, acropodite wanting........................... ............................................................................... Doratodesmus hispidus

15 Body with 20 segments. Metatergal tuberculation less distinct, mostly like three transverse rows of bosses ( Figs 26 A-C). Solenomere branch very short, acropodite rather short and simple ( Figs 29B, C ). China .............................. ...................................................................... Doratodesmus grandifoliatus

– Body with 19 segments. Metatergal tubercles also mostly arranged in three transverse rows, but very distinct, conical. Solenomere branch absent, acropodite very long, falcate and conspicuously spinose. Japan............................ ................................................................................. Doratodesmus elegans

16 Body with 20 segments. At least some metaterga with an evident mid-dorsal outgrowth or projection ( Figs 5A, C )........................................................ 17

– Body with 19 or 20 segments. All metaterga subequal, devoid of an evident mid-dorsal outgrowth or projection ( Figs 1A ; 22A) .................................. 22

17 Only last 3-8 metaterga in front of telson with an evident mid-dorsal outgrowth ( Figs 5A, C ) .................................................................................. 18

– Most metaterga, including some of anterior body portion, with a high, often tuberculated projection............................................................................. 20

18 Metaterga 12-19 each with an increasingly evident, subtriangular, mid-dorsal outgrowth. Cave in Yunnan Province, China ... Eutrichodesmus dorsiangulatus

– Only metaterga 16(17)–19 each with an evident, rather rounded, mid-dorsal outgrowth................................................................................................. 19

19 Paraterga narrower ( Fig. 6B ). Gonopod process dp much shorter ( Figs 7 ; 8 ). North Vietnam .............................. Eutrichodesmus armatocaudatus sp. n.

– Paraterga broader. Gonopod process dp much longer. Cave in Yunnan Province, China ..................................................... Eutrichodesmus monodentus

20 Mid-dorsal projections increasingly evident on metaterga 3-18, abruptly smaller on metatergum 19. Distal half of gonopod telopodite enlarged, lobuliform, fringed apically..................................... Eutrichodesmus cavernicola

– Mid-dorsal projections especially prominent, present on metaterga 5-19, only slightly less prominent on metatergum 19. Male, when known ( E. macclurei ), with distal half of gonopod telopodite slender........................................... 21

21 Mid-dorsal projections on metaterga 5 and 6 straight in lateral view............. ........................................................................... Eutrichodesmus macclurei

– Mid-dorsal projections on metaterga 5 and 6 slightly inclined anteriorly in lateral view......................................................... Eutrichodesmus reclinatus

22 Body with 19 segments. Collum and metaterga very densely setose, setae long, bisegmented and tactile ( Figs 9 ; 12 A-C). Gonopod telopodite with a highly peculiar dp ( Fig. 13 )................ Eutrichodesmus communicans sp. n.

– Body with 20 segments. Collum and metaterga without dense setation. Gonopod telopodite with a different dp......................................................... 23

23 Most paraterga very wide and only slightly declivous ( Figs 14 A-D; 15C; 18; 19D). Caves in Guangxi Province, China ................................................. 24

– Paraterga not so wide, strongly declivous ( Figs 1 ; 2D; 22C; 23E) ............. 25

24 Collum with a conspicuous row of teeth all along fore margin of collum ( Figs 18D ; 19A ). Gonopods as in Figs 20C, D ; 21 ... Eutrichodesmus similis sp. n.

– Collum without such conspicuous teeth at fore margin of collum ( Figs 15A, B ). Gonopods as in Figs 16 B-E; 17B, C............ Eutrichodesmus latus sp. n.

25 Most metaterga with two transverse rows of bosses. Epiproct very strongly flattened dorsoventrally, subquadrate-spatuliform, with unincised margins. Gonopod telopodite particularly slender, about twice as long as coxa. Japan........ ........................................................................... Eutrichodesmus peculiaris

– Most metaterga with three transverse rows of bosses or conical tubercles. Epiproct never strongly flattened. Gonopods telopodite shorter..................... 26

26 Paraterga mostly set off laterally at base by a deep sulcus, metaterga with conical tubercles. Gonopod telopodite with a particularly small, dentiform dp.... ............................................................................ Eutrichodesmus gremialis

– Paraterga not set off laterally at base, continuing general outline of metaterga. Gonopod telopodite with a larger and longer dp....................................... 27

27 Body up to 5.0 mm long. Paraterga relatively narrow ( Figs 1 ; 2D), volvation imperfect. Gonopods relatively simple ( Figs 4B, C ) ..... Eutrichodesmus basalis sp. n.

– Body at least 7.0 mm long. Paraterga broader ( Figs 22 ; 23E ), volvation complete. Gonopods more elaborate ............................................................... 28

28 Caudolateral and lateral lobulations on most of metaterga deeply incised ( Fig. 22 ). Gonopods with a peculiar, long, biramous, multituberculate dp ( Figs 25 ; 26B, C ).............................................................. Eutrichodesmus incisus sp. n.

– Metaterga not incised caudolaterally. Process dp on gonopods shorter, simple, unciform and pointed ............................................................................... 29

29 Body ca 8.0 mm long. Gonopod tip bifid. Vietnam...................................... ........................................................................... Eutrichodesmus demangei

– Body ca 14.0 mm long. Gonopod tip simple, unciform. Cave in Yunnan Province, China ............................................... Eutrichodesmus arcicollaris

Figures

Figure 1. Eutrichodesmus basalis sp. n., ♀ paratype; A, habitus, lateral view; B, anterior part of body, lateral view; C, midbody segments, lateral view; D, posterior part of body, lateral view; E, midbody segments, dorsal view; F, posterior part of body, dorsal view. − Scale bars: A, 0.5 mm, B, C, D & F, 0.1 mm & E, 0.2 mm.

Figure 4. Eutrichodesmus basalis sp. n., ♂ paratype; A, leg 9, caudal view; B & C, left gonopod, mesal and lateral views, respectively (dp = distofemoral process). − Scale bar: A, 0.2 mm, B, C, 0.1 mm.

Figure 5. Eutrichodesmus armatocaudatus sp. n., ♂ paratype from cave 4; A, habitus, lateral view; B, midbody segments, lateral view; C, posterior part of body, lateral view; D, midbody segments, ventral view; E, posterior part of body, ventral view; F, telson, ventral view. − Scale bars: A, 1.0 mm, B, C, E, 0.2 mm, D, 0.5 mm & F, 0.1 mm.

Figure 6. Eutrichodesmus armatocaudatus sp. n., ♂ paratype from cave 4; A, anterior part of body, frontoventral view; B, cross-section of a midbody segment, caudal view; C, enlarged paraterga with ozopores, lateral view; D, paratergum with porostele, caudal view; E, porostele; F, texture of prozonite and limbus; G, cerotegument with a bisegmented seta. − Scale bars: A, B, 0.2 mm, C, D, 0.1 mm, E, G, 0.01 mm & F, 0.05 mm.

Figure 7. Eutrichodesmus armatocaudatus sp. n., ♂ paratype from cave 4; A, body segments 6 and 7, ventral view; B, gonopods, ventral view; C, same, ventral view; D, gonopod apices. − Scale bars: A, 0.5 mm, B, 0.2 mm, C, 0.05 mm & D, 0.02 mm.

Figure 8. Eutrichodesmus armatocaudatus sp. n., ♂ paratype from cave 4; A & B, right gonopod, mesal and lateral views, respectively (micropilosity not shown; dp = distofemoral process). − Scale bar: 0.2 mm.

Figure 9. Eutrichodesmus communicans sp. n., ♂ paratype from Malo Island, habitus.

Figure 12. Eutrichodesmus communicans sp. n., ♂ paratype from Fapon doline 3; A, metatergal cerotegument and bisegmented setae; B, same with microvilli, lateral view; C, same with base of a bisegmented seta; D, legs, ventral view; E, gonopods, ventral view; F, left gonopod, dorsolateral view. − Scale bars: A, B, 0.01 mm, C, 0.005 mm, D, F, 0.05 mm.

Figure 13. Eutrichodesmus communicans sp. n., ♂ paratype from Boutmas; A-D, left (A, B) and right (C, D) gonopods, lateral, mesal, mesal and lateral views, respectively (dp = distofemoral outgrowth). − Scale bar: 0.2 mm.

Figure 14. Eutrichodesmus latus sp. n., ♂ paratype from Yan Wu Dong Cave; A, habitus, lateral view; B, anterior part of body, lateral view; C, posterior part of body, lateral view; D, same, dorsal view; E, same, ventral view; F, anterior part of body, ventral view. − Scale bars: A, 1.0 mm, B, C, E, 0.2 mm, D, F, 0.5 mm.

Figure 15. Eutrichodesmus latus sp. n., ♂ paratype from Yan Wu Dong Cave; A & B, anterior part of body, front and dorsal views, respectively; C, cross-section of segment 6, caudal view; D, midbody paratergum with ozopore; E, texture of prozonite and limbus; F, metatergal cerotegument with a bisegmented seta. − Scale bars: A, C, 0.5 mm, B, 0.2 mm, D, 0.1 mm, E, 0.02 mm & F, 0.01 mm.

Figure 16. Eutrichodesmus latus sp. n., ♂ paratype from Yan Wu Dong Cave; A, midbody sterna, ventral view; B, gonopods, ventral view; C, same, ventral view; D, gonopod apices, ventral view; E, tip of distofemoral process of gonopods, ventral view. − Scale bars: A-C, 0.1 mm, D & E, 0.02 mm.

Figure 18. Eutrichodesmus similis sp. n., ♂ paratype from Gui Dong 2 Cave; 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 19. Eutrichodesmus similis sp. n., ♂ paratype from Gui Dong 2 Cave; A, anterior part of body, ventral view; B, segments 5-9, ventral view; C, posterior part of body, ventral view; D, cross-section of segment 4, caudal view; E, texture of prozonite, limbus and bisegmented seta; F, metatergal cerotegument with a bisegmented seta. − Scale bars: A, D, 0.5 mm, B, 1.0 mm, C, 0.02, E, 0.01 mm & F, 0.005 mm.

Figure 20. Eutrichodesmus similis sp. n., ♂ paratype from Gui Dong 2 Cave; A, paratergum with ozopore, lateral view; B, leg; C, gonopods, ventral view; D, gonopod apex, ventral view. − Scale bars: A-C, 0.1 mm & D, 0.05 mm.

Figure 21. Eutrichodesmus similis sp. n., ♂ paratype from Gui Dong 2 Cave; A, left gonopod, mesal view; B & C, distal half of right gonopod, mesal and lateral views, respectively (dp = distofemoral process). − Scale bar: 0.2 mm.

Figure 22. Eutrichodesmus incisus sp. n., ♀ paratype from Tiao Shuz Dong Cave; A, habitus, lateral view; B, anterior part of body, sublateral view; C, midbody segments, lateral view; D, posterior part of body, lateral view; E, anterior part of body, frontodorsal view; F, midbody segments, dorsal view. − Scale bars: A, 0.5 mm & B-F, 0.2 mm.

Figure 23. Eutrichodesmus incisus sp. n., ♀ paratype from Tiao Shuz Dong Cave; A, posterior part of body, dorsal view; B, anterior part of body, subventral view; C, midbody segments, ventral view; D, telson, ventral view; E, cross-section of a midbody segment, caudal view; F, paratergum with ozopore, lateral view. − Scale bars: A-C, E, 0.2 mm, D, 0.1 mm & F, 0.05 mm.

Figure 25. Eutrichodesmus incisus sp. n., ♂ paratype from Liao Jing Ling Cave; A, gonopods, ventral view; B, distal parts of gonopods, ventral. − Scale bars: A, 0.1 mm & B, 0.05 mm.

Figure 26. Eutrichodesmus incisus sp. n., ♂ paratype from Tiao Shuz Dong Cave; A, leg 9, caudal view; B & C, right gonopod, mesal and lateral views, respectively (dp = distofemoral process). − Scale bar: A, 0.2 mm, B, C, 0.1 mm.

Figure 29. Doratodesmus grandifoliatus (Zhang in Zhang & Wang, 1993), ♀ from Ma Fa Tiao Dong Cave; A, texture of pro- and metazona; B, bisegmented seta; C, leg and sternum, caudal view; D, claw. − Scale bars: A, 0.05 mm, B, 0.002 mm, C, 0.1 mm & D, 0.02 mm.