Corresponding author: Sergei I. Golovatch (
Academic editor: R. Mesibov
The
Golovatch SI, Geoffroy J-J, Mauriès J-P, VandenSpiegel D (2015) Review of the millipede genus
The millipede family
The present paper puts on record another nine new species of
The material serving as the basis for the present contribution derives from subterranean collections made in China by Josiane Lips (Villeurbanne, France), Leonardo Latella and Daniele Avesani (both from the Museo Civico di Storia naturale, Verona, Italy), as well as Boris Sket, Peter Trontelj and their collaborators (all from the University of Ljubljana, Slovenia). All material, including the holotypes, has been deposited in MNHN. The term “doratodesmoid” is used hereafter only in its vernacular meaning, in order to concisely characterize a body shape, i.e. capable or nearly capable of volvation.
SEM micrographs were taken using a JEOL JSM-6480LV scanning electron microscope.
After examination, SEM material was removed from stubs and returned to alcohol, all such samples being kept at MNHN.
Holotype ♂ (MNHN JC 367), China, Sichuan Prov., Beichuan County, Cave Yan Dong, 18.VIII.2004, leg. J. Lips (No. 1583).
To emphasize the prominent, triangular, distofemoral process (
Differs from congeners by the prominent, triangular, distofemoral process of the gonopod (see also Key below).
Length ca 7.0 mm, width 0.9 and 1.5 mm on midbody pro- and metazonae, respectively. Coloration uniformly very light brown with pallid antennae, clypeolabral region, prozonae, venter, legs and metatergal tuberculations (Fig.
Body with 19 segments (♂) (Fig.
Sterna usually with a deep and narrow depression between coxae. Legs short, crassate except for slender tarsi, about as long as body height.
Gonopods (Fig.
The presence of only 19 body segments is rare in
More information on the location of the cave can be found at
Holotype ♂ (MNHN JC 368), China, Guangxi Prov., Guilin County, Grotte des Squelettes, 22.VII.1992, leg. J. Lips (No. B1-2).
Paratypes: 1 ♂ (SEM), 1 juv. (MNHN JC 368), same data, together with holotype.
In honour of Josiane Lips, the collector.
Differs from congeners by clearly elevated mid-dorsal regions of most metaterga, coupled with a slender, suberect gonopod telopodite which shows a rather narrowly gapped apical pincer (see also Key below).
Length of adults ca 7.0 mm, width 1.0 and 1.95 mm on midbody pro- and metazonae, respectively (♂). Coloration entirely pallid.
All characters as in
Body with 20 segments (♂) (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Figs
More information on the location of the cave can be found at
Holotype ♂ (MNHN JC 369), China, Guizhou Prov., Guanling County, Yong Ning Town, Cave Yun Dong (Cloud Cave), 01.VIII.2005, leg. L. Latella & D. Avesani.
Paratypes: 1 ♀, 2 subadult ♀ (MNHN JC 369), 1 subadult ♀ (SEM), same data, together with holotype.
To emphasize the relatively slender body due to subvertical paraterga; adjective.
Differs from congeners by the large body size, clearly elevated mid-dorsal regions of most metaterga, coupled with narrow, strongly declivous, subvertical paraterga and a simple, falcate gonopod telopodite carrying a long, spiniform, distofemoral process (see also Key below).
Length of adults ca 14–15 mm, width 1.8 and 2.5 mm (♂ holotype) or 1.6 and 2.1 mm (♀ paratype) on midbody pro- and metazonae, respectively. Coloration entirely pallid, sometimes (♀ paratype) with traces of reddish earth material on metaterga.
All characters as in
Body with 20 segments (♂, ♀), conglobation pattern typical of “doratodesmoids”, volvation apparently being complete because of particularly strongly declivous and short paraterga. Antennae short and clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Fig.
More information on this cave and its fauna can be found in
Holotype ♂ (MNHN JC 370), China, Guizhou Prov., Libo County, Libo, Cave Feng Dong, 07.III.1995, leg. P. Trontelj.
Paratypes: 1 ♂, 1 juv. ♀ (MNHN JC 370), 1 ♂ (SEM), same data, together with holotype.
Non-types: 1 ♂, 2 ♀, 11 subadult ♀ or juv. (MNHN JC 370), 1 ♂ (SEM), China, Guizhou Prov., Libo County, Shuipa, Cave Shui Jiang Dong – Cave Shuipu Da Dong, 28.II.1995, leg. P. Trontelj; 1 ♂ (MNHN JC 370), 1 ♂ (SEM), Guizhou Prov., Libo County, Jia Ban, Cave La Tai Dong, 06.III.1995, leg. P. Trontelj.
In honour of Peter Trontelj, the collector.
Differs from congeners by the particularly broad and moderately declivous paraterga which are set at about 45° to the vertical axis and continue the outline of the sides above paraterga, coupled with mostly 4-5 irregular rows of flat setigerous tubercles/bosses per metatergum, the calyx-shaped tergal setae, and the fairly complex gonopod (see also Key below).
Length of adults ca 8-9 mm, width 1.2 and 2.2 mm (♂ paratype and one ♀ non-type from Shui Jiang Dong) to 1.5 and 2.5 mm (♂ holotype and other non-types) on midbody pro- and metazonae, respectively. Coloration entirely pallid, except some traces of reddish earth material on paraterga.
All characters as in
Body with 20 segments (♂, ♀), conglobation pattern typical of “doratodesmoids”, volvation apparently being incomplete because of particularly broad and only moderately declivous paraterga. Antennae rather long and poorly clavate (Figs
Sterna usually with a rather deep, narrow depression between coxae (Figs
Gonopods (Figs
The conspecificity of the non-type samples with
Interestingly, calyx-shaped tergal setae among
Holotype ♂ (MNHN JC 371), China, Guizhou Prov., Zhen Feng County, Bei Pan Jiang Town, Cave Shui Chi Dong (Water Pool Cave), ca. 1060 m a.s.l., 31.VII.2005, leg. L. Latella & D. Avesani.
Paratypes: 1 ♂, 2 ♀ (MNHN JC 371), 1 ♀ (SEM), same data, together with holotype.
In honour of Leonardo Latella, one of the main collectors.
Differs from congeners by the broad and moderately declivous paraterga which are set at about 45° to the vertical axis and almost continue the outline of the sides above paraterga, coupled with three irregular rows of flat setigerous bosses per metatergum, and the especially simple gonopod (see also Key below).
Length of adults ca 12–13 mm, width 1.1–1.2 and 2.8–3.0 mm on midbody pro- and metazonae, respectively (♂, ♀). Holotype ca 12 mm long, 1.2 and 3.0 mm wide on midbody pro- and metazonae, respectively. Coloration entirely pallid, except some traces of reddish earth material on terga.
All characters as in
Body with 20 segments (♂, ♀), conglobation pattern typical of “doratodesmoids”, volvation apparently being incomplete because of particularly broad and only moderately declivous paraterga. Antennae rather long and poorly clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Fig.
More information on this cave and its fauna can be found in
Holotype ♂ (MNHN JC 372), China, Guizhou Prov., Guanling County, Huajiang Town, Cave Huashiban Dong (Slippery Cave), 26.VII.2005, leg. L. Latella & D. Avesani.
Paratypes: 1 ♂ (MNHN JC 372), 1 ♀ (SEM), same data, together with holotype.
To emphasize the mostly obliterate metatergal tuberculation; adjective.
Differs from congeners by the largely obliterate metatergal tuberculation (even those at the fore margin of metetergum 2), the rather broad and strongly declivous paraterga which are set at about 30° to the vertical axis and continue the outline of the sides above paraterga, coupled with three irregular rows of very flat setigerous bosses per metatergum, and the fairly complex gonopod telopodite (see also Key below).
Length of adults ca 10 mm, width 1.0–1.1 and 2.1–2.2 mm on midbody pro- and metazonae, respectively (♂, ♀). Holotype ca 1.1 and 2.2 mm wide on midbody pro- and metazonae, respectively. Coloration entirely pallid.
All characters as in
Body with 20 segments (♂, ♀), conglobation pattern typical of “doratodesmoids”, volvation apparently being incomplete because of broad and only rather strongly declivous paraterga. Antennae rather long and poorly clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Fig.
More information on this cave and its fauna can be found in
Holotype ♂ (MNHN JC 373), China, Hunan Prov., Longshan County, Huaoyan, Cave Feihu Dong (33a), 13.IV.1997, leg. B. Sket, Cao & R. Verovnik.
Paratype: 1 subadult ♀ (SEM), same data, together with holotype.
In honour of Boris Sket, one of the main collectors.
Differs from congeners by the relatively narrow and strongly declivous paraterga which are set low on the body at about 30° to the vertical axis and nearly continue the outline of the sides above paraterga, the low, but distinct, rounded, metatergal bosses arranged in three transverse irregular rows, and the rather simple gonopod (see also Key below).
Length of holotype ca 7 mm, width 1.0 and 1.7 mm on midbody pro- and metazonae, respectively. Coloration entirely pallid.
All characters as in
Body with 20 segments (♂), conglobation pattern typical of “doratodesmoids”, volvation apparently being complete because of narrow and strongly declivous paraterga. Antennae rather short and clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Fig.
Holotype ♂ (MNHN JC 374), China, Hubei Prov., Yishang Yichang County, Grotte des Araignées, 15.VIII.1992, leg. J. Lips (K1-2).
Paratype: 1 ♂ (SEM), same data, together with holotype.
To emphasize the apical termination of the seminal groove; adjective.
Differs from congeners by the relatively narrow and strongly declivous paraterga which are set low on the body at about 40° to the vertical axis and distinctly discontinue the subvertical outline of the sides above paraterga, coupled with narrow paraterga which only slightly overreach the level of the venter, the low, but distinct, rounded, metatergal tuberculations arranged in three transverse irregular rows, and the rather complex gonopod (see also Key below).
Length of holotype ca 7 mm, width 0.6 and 1.0 mm on midbody pro- and metazonae, respectively. Coloration entirely pallid.
All characters as in
Body with 20 segments (♂), conglobation pattern typical of “doratodesmoids”, volvation apparently being complete because of narrow and strongly declivous paraterga. Antennae rather short and clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Figs
More information on the location of the cave can be found at
Holotype ♂ (MNHN JC 375), China, Guizhou Prov., Kaiyang, Cave Xianyan Dong, 19.II.2004, leg. S. Prevorčnik & B. Sket.
Paratypes: 1 ♂, 2 ♀, 2 subadult ♀ (MNHN JC 375), 1 ♀ (SEM), same data, together with holotype.
To emphasize cavernicoly; adjective.
Differs from congeners by the relatively broad and modestly declivous paraterga which are set low on the body at about 45° to the vertical axis and distinctly discontinue the more strongly declined outline of the sides above paraterga, coupled with low, but distinct, mostly longitudinally oblong, metatergal tuberculations arranged in three transverse irregular rows, and the rather simple gonopod acropodite which only shows a small distodorsal tooth (see also Key below).
Length of adults ca 8–9 mm, width 1.0 and 1.9–2.0 mm on midbody pro- and metazonae, respectively. Holotype ca 9 mm long, 1.0 and 2.0 mm wide on pro- and metazonae, respectiverly. Coloration entirely pallid.
All characters as in
Body with 20 segments (♂, ♀), conglobation pattern typical of “doratodesmoids”, volvation apparently being incomplete because of broad and modestly declivous paraterga. Antennae rather long and poorly clavate (Fig.
Sterna usually with a rather deep, narrow depression between coxae (Fig.
Gonopods (Fig.
The nine new species described here are presumed to be troglobites, as all were collected in caves and all are troglonorphic as evidenced by unpigmented teguments. Their discovery supports the ideas that
1 | Each postcollum metatergum with only two transverse rows of tuberculations or bosses |
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– | Each postcollum metatergum with at least three transverse rows of tuberculations or bosses |
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2 | 1+1 mid-dorsal tubercles only slightly higher than others and located only in 2nd row on segments 4-6(7). Zhejiang Province |
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– | Mid-dorsal tubercles much higher than others and located in both rows at least on segments 4-16(17) |
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3 | Most of tuberculations on collum obliterated, retained only near lateral edge. Mid-dorsal tubercles on penultimate segment low, but evident, like a small crest. Chongqing Municipality |
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– | Almost entire collum covered with tuberculations. Mid-dorsal tubercles on penultimate segment nearly wanting, flat, not crest-shaped. Fujian Province |
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4 | Adult body with 19 segments (Fig. |
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– | Adult body with 20 segments. Distofemoral process of gonopod not triangular and acuminate |
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5 | At least some metaterga increasingly strongly elevated mid-dorsally towards segment 17 or 18, with 1-2 outgrowths, projections or a ridge (e.g. Figs |
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– | No metaterga elevated mid-dorsally over others. Three or more transverse rows of tuberculations or bosses per metatergum |
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6 | Mid-dorsal regions of metaterga increasingly strongly elevated towards segment 17 due to enlarged tubercles of middle row, thereafter smaller (Fig. |
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– | Mid-dorsal regions of metaterga increasingly strongly elevated towards segment 18. Gonopod distofemoral process held subrectangular to acropodite, gap between |
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7 | Metaterga 7-18 each with an increasingly evident mid-dorsal outgrowth/crest, bimodal due to clearly enlarged 1st and 2nd rows of tuberculations. Distofemoral process of gonopod microtuberculate |
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– | Mid-dorsal crests on metaterga unimodal, subtriangular due to only 2nd row of tuberculations being clearly enlarged. Distofemoral process of gonopod a simple long spine |
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8 | Paraterga narrow, subvertical (Fig. |
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– | Paraterga considerably broader, directed ventrolaterad. Gonopod acropodite only slightly curved ventrad, devoid of a hairpad. Yunnan |
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9 | Metatergal tuberculations very small knobs. Gonopod acropodite clearly enlarged relative to distofemoral process, devoid of a distodorsal tooth |
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– | Metatergal tuberculations mostly distinct. Gonopod acropodite slender, with a distodorsal tooth |
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10 | At least some metaterga with tuberculations/bosses arranged in 4-5 transverse irregular rows |
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– | All metaterga with only three transverse rows of tuberculations or bosses |
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11 | Metaterga 2-13 each with four, following ones with five, rows of tuberculations or bosses,. Guangdong |
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– | Metaterga with 3-5 rows of tuberculations or bosses, pattern of increase different |
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12 | Each postcollum metatergum with 4-5 irregular rows of bosses (Figs |
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– | Metaterga with 3-4 rows of tuberculations or bosses. Gonopod distofemoral process ( |
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13 | Paraterga narrow, set low on body, rather strongly declined ventrolaterad and more or less clearly discontinuing the outline of sides above paraterga |
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– | Paraterga broad to very broad, set higher on body to (almost) continue the outline of sides above paraterga |
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14 | Distofemoral process of gonopod a simple, strong, ventrobasally setose hook directed dorsad. Yunnan |
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– | Distofemoral process of gonopod more elaborate |
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15 | Seminal groove terminating apically, distofemoral process ( |
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– | Seminal groove terminating subapically, distofemoral process of gonopod devoid of micropilosity |
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16 | Distofemoral process of gonopod bipartite, long and complex. Guizhou |
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– | Distofemoral process ( |
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17 | Paraterga very broad, each about as wide as prozonite. Guangxi |
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– | Paraterga considerably narrower than prozonite width | 19 |
18 | Collum devoid of a row of lobulations at fore margin |
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– | Collum with a row of distinct lobulations at fore margin |
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19 | Gonopod simple, but unusually strongly falcate. Hunan |
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– | Gonopod only slightly to moderately curved |
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20 | Distofemoral process of gonopod strongly appressed to a simple acropodite. Jiangxi |
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– | Distofemoral process of gonopod not appressed to often a more elaborate acropodite, gap between both parts being considerable. Guizhou |
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21 | Gonopod acropodite complex, with a number of teeth ( |
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– | Gonopod acropodite simple, at most with a small tooth |
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22 | Gonopod acropodite with a small tooth |
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– | Gonopod acropodite devoid of considerable outgrowths (Fig. |
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This work only became possible through the support provided to the first author by the MNHN. We are most grateful to all collectors who entrusted us their material for study and allowed to deposit it in MNHN. Alexandr Kirejtshuk (St. Petersburg, Russia) skillfully took pictures in Fig.