Riotintobolus mandenensis Wesener, 2009, by original designation.

Riotintobolus minutus Wesener, 2009

Riotintobolus aridus Wesener, 2009

Riotintobolus anomalus Wesener, 2009

Riotintobolus tsimelahy sp. nov.

Riotintobolus mangatsiaka sp. nov.

Riotintobolus lavanono sp. nov.

Riotintobolus bovinus sp. nov.

Riotintobolus antafoky sp. nov.

Riotintobolus makayi sp. nov.

Southern spiny forests and gallery forests, southeastern littoral rainforests (Fig. 2).

Figure 2. 

Distribution of the newly described Riotintobolus species, as well as of the still undescribed species. Vegetation marks modified from Moat and Smith 2007.

1 ♂ holotype, ZFMK MYR9940, Madagascar, PN Andohahela, Tsimelahy, 24°57.296'S, 046°37.214'E, 135 m, spiny forest, coll. Wesener and Schütte, 24.v.2007.

Paratypes : 9 ♂, 12 ♀, ZFMK MYR941, same data as holotype; 1 ♂, ZFMK MYR9950, same data as holotype; 1 ♂ with damaged gonopods; ZFMK MYR9949, same data as holotype.

3 ♂ and ♀, BLF 5239 (CASENT9032808), Madagascar, Toliara, Forêt de Mahavelo, Isantoria River, 24°45'30"S, 46°9'26"E, 110 m, 28.i-1.ii.2002, spiny forest/thicket, EH18 pitfall trap, coll. B. L. Fisher et al.; 1 ♂, 1 imm., MZUF Fi-10, Madagascar; Andohahela, 6–12.xii.1991, leg B. Randriamampionona.

Tsimelahy, after the type locality (Fig. 2), spiny forests next to the Tsimelahy River, Andohahela National Parc. Noun in apposition.

Riotintobolus tsimelahy sp. nov. shares the absence of a projecting epiproct on the telson only with R. anomalus, R. antafoky sp. nov., R. bovinus sp. nov., R. mangatsiaka sp. nov. and R. lavanono sp. nov. The posterior telopod featuring two slender, sharp projections is only shared with R. bovinus sp. nov., R. mangatsiaka sp. nov. and R. lavanono sp. nov. A posterior gonopod separated into three parts is only shared with R. mangatsiaka sp. nov. and R. lavanono sp. nov., whose habitus and gonopods look very similar to those of R. tsimelahy sp. nov. Both species differ in details of the tip of the posterior gonopod and in the colour of their antennae and legs, which are red in R. tsimelahy sp. nov. and dark grey in R. lavanono sp. nov. R. tsimelahy sp. nov. differs from R. mangatsiaka sp. nov. in the presence of two lateral processes on the posterior gonopod. All three species differ by >11% uncorrected p-distance in the COI barcoding gene.

Measurements : Telson not included in counts of segments. Male holotype with 50+0 segments, ca. 44 mm long, 4.3 mm wide. Largest females of type series with 49 or 50+0 segments, up to 50 mm long, 5.3 mm wide.

Colour (in living specimens): Body rings grey, appendages red. Head, paraprocts and posterior margins of body segments darker grey to black. Ozopore openings highlighted by black spot (Fig. 3A, C).

Head : each eye with 30–35 ommatidia in six rows. Incisura lateralis open (Fig. 3A). Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side (Fig. 3B). Antennae long, protruding back to segment 5. Length of antennomeres: 1<2>3=4=5=6. Second antennomere slenderer but twice as long as first. Terminal antennomere with four large sensory cones located together inside a membranous area.

Figure 3. 

Riotintobolus tsimelahy sp. nov., holotype (ZFMK MYR99940) (A–F) and male paratype (ZFMK MYR9949) (G, H) A multi-layer photograph, male anterior body, lateral view B multi-layer photograph, male leg pairs 1–7, ventral view C multi-layer photograph, male posterior body end with telson D anterior gonopod, anterior view E left posterior gonopod, posterior view F right posterior gonopod, anterior view G damaged? left posterior gonopod, posterior view H damaged? right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; z = apical membranous ‘flag’; numbers refer to leg pair number. Scale bars: 1 mm.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Palpi of similar size. Endochilarium not dissected.

Mandible not dissected.

Collum : smooth, laterally not protruding as far as ring 2 (Fig. 3A).

Body rings : ozopores starting at segment 6, marked by a black spot. Located on suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts elongated, with weak lips, abundant micropunctation especially towards edges (Fig. 3C). Epiproct well-rounded, covering, but not reaching above paraproct (Fig. 3C). Hypoproct inconspicuous (Fig. 3C).

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa. Tarsus with three pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus with a single apical and three pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in males.

Female sexual characters. No tarsal pads, antennae shorter than male, only protruding back to ring 2. Vulvae not dissected.

Male sexual characters : tarsal pads present from leg 3 to midbody, small, inconspicuous (Fig. 3B). Coxae 3–7 without coxal processes, but coxae 3–5 swollen (Fig. 3B).

Anterior gonopod sternite massive (Fig. 3D), elongated into a wide, well-rounded triangular lobe (Fig. 3D). Sternite in anterior view well-visible, without discernible apodemes, protruding almost higher than coxal processes. Coxite with a large, well-rounded mesal process (Fig. 3D). Telopodite with process arising mesally (Fig. 3D), process apically curved with a large triangular projection (Fig. 3D), tip well-rounded, slightly protruding above lateral margin of telopodite (Fig. 3D). Whole telopodite process resembling an even-sided triangle.

Posterior gonopods consisting of three parts, separated by sutures or articulations: a basal coxite with a slender coxite projection and a slightly shorter telopodite, efferent duct discharging laterally (Fig. 3E, F). Process of coxite and telopodite standing in same axis (Fig. 3E, F). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite. Basal part of coxite wide, mesally with a large triangular sclerite located on lower level than remaining part. Coxite elongated. Efferent duct running at mesal margin of coxite (Fig. 3E) before curving to the lateral margin at beginning of telopodite (Fig. 3E). Telopodite as wide as but much shorter than coxite, standing in same axis (Fig. 3F), apically membranous, with one triangular apical process and two slender lateral processes (Fig. 3E, F). Lateral processes straight, running almost parallel to one another, slender and sclerotised, efferent duct seems to be ending at base of lateral process (Fig. 3E). Base of lateral process with a short, membranous-white projection (Fig. 3E).

The number of segments varies, even within one population, between 47 and 51. The population from the Forêt de Mahavelo (Fig. 1) differs by 1.5% uncorrected p-distance of the COI gene to those from the type locality, Tsimelahy (Suppl. material 1). One small male from the type series (ZFMK MYR9949) has incompletely developed posterior gonopods, either a sign of healed damage (there are black spots on it), or maybe not fully developed (FiIg. 3G, H).

R. tsimelahy sp. nov. could be found in great numbers in the early morning (7–9 a.m.) on the forest floor of the spiny bush. The otherwise dry spiny bush was still quite wet because of dew. No juveniles were observed. Contrary to other Riotintobolus species, such as R. mandenensis and R. minutus, R. tsimelahy sp. nov. did not remain stiff like a stick when disturbed, but rolled-up into a spiral, a common defence behaviour for juliform millipedes.

1 ♂ holotype, ZFMK MYR9801, Madagascar, PN Andohahela, Mangatsiaka, 24°58.051'S, 046°33.206'E, 90 m, spiny forest, leg. Wesener and Schütte, 23.v.2007.

Paratypes : 7 ♂, 14 ♀, ZFMK MYR938, same data as holotype.

1 ♂, 1 ♀, FMMC 5413, Province Toliara; RNI d’Andohahela, parcel 2; 120 m; 24°49.0'S, 46°36.6'E; pitfalls, camp 6; leg. S. Goodman 7–15.xii.1995; 1 ♂, FMMC 5379, Province Toliara; RNI d’Andohahela, parcel 2; 120 m; 24°49.0'S, 46°36.6'E; pitfalls, camp 6; leg. S. Goodman 7–15.xii.1995;

Mangatsiaka, after the type locality (Fig. 2), spiny forests next to a site called Mangatsiaka, Andohahela National Parc. Noun in apposition.

Riotintobolus mangatsiaka sp. nov. shares the absence of a projecting epiproct on the telson only with R. anomalus, R. antafoky sp. nov., R. bovinus sp. nov., R. tsimelahy sp. nov. and R. lavanono sp. nov. The posterior telopod featuring two slender, sharp projections is only shared with R. bovinus sp. nov., R. tsimelahy sp. nov. and R. lavanono sp. nov. A posterior gonopod separated into three parts is only shared with R. tsimelahy sp. nov. and R. lavanono sp. nov., whose habitus and gonopods look very similar to those of R. mangatsiaka sp. nov. Both species differ in details of the tip of the posterior gonopod and in the colour of their antennae and legs, which are red in R. mangatsiaka sp. nov. and dark grey in R. lavanono sp. nov. R. mangatsiaka sp. nov. differs from R. tsimelahy sp. nov. in the presence of just one lateral processes on the posterior gonopod. All three species differ by >11% uncorrected p-distance in the COI barcoding gene.

Measurements : male holotype with 49+0 segments, ca. 42 mm long, 4.1 mm wide. Largest females of type series with 48 to 51+0 segments, up to 52 mm long, 5.4 mm wide.

Colour (in living specimens): Body rings grey, appendages red. Head, paraprocts and posterior margins of body segments darker grey to black (Fig. 4A). Ozopore openings highlighted by black spot (Fig. 4A, B).

Figure 4. 

Riotintobolus mangatsiaka sp. nov., paratypes (ZFMK MYR938) A living female B living male in defensive position C multi-layer photograph, male anterior body, lateral view D multi-layer photograph, male leg pairs 1–7, ventral view E multi-layer photograph, male posterior body end with telson F anterior gonopod, anterior view G anterior gonopod, posterior view H right posterior gonopod, posterior view I right posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Md = basal joints of mandible; Pe = penis; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; z = apical membranous ‘flag’; numbers refer to leg pair number. Scale bars: 1 mm.

Head : each eye with 30–35 ommatidia in six rows. Incisura lateralis open (Fig. 4C). Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side (Fig. 4D). Antennae long, protruding back to segment 5. Length of antennomeres: 1<2>3=4=5=6. Second antennomere slenderer but twice as long as first. Terminal antennomere with four large sensory cones located together inside a membranous area (Fig. 5A). Antennomere 5 with 3 rows, antennomere 6 latero-apically with a single row of sensilla basiconica (Fig. 5B–E). Antennomere 6 with an unknown type of at least three sensilla or duct openings located close to disc (Fig. 5D, E).

Figure 5. 

Riotintobolus mangatsiaka sp. nov., male (FMMC 5413), SEM A left antennae B antennomere 5 C details of sensilla basiconica D antennomere 6 E detail of unknown sensilla/pore opening F gnathochilarium underside G gnathochilarium, detail of right central pad. Abbreviations: cP = central pads; d = disc; iP = inner palpus; LP = lateral palpus; s = unknown sensillum; sb = sensilla basiconica; numbers refer to antennomere number. Scale bars as indicated.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Palpi of similar size (Fig. 5F). Central pads with standard two sets of sensory cones, apical area with ten cones, higher area with ~ 30 (Fig. 5G).

Mandible : Stipes without projection, well rounded (Fig. 4C). Gnathal lobe, external tooth simple, rounded; mesal tooth with three cusps (Fig. 6A). Eight pectinate lamellae. Mesal margin of pectinate area (intermediate area) with ca. four rows of small, slender spines. Molar plate with few, five, transverse furrows (Fig. 6A).

Figure 6. 

Riotintobolus mangatsiaka sp. nov., male and female (FMMC 5413), SEM A left gnathal lobe of mandible, mesal view B left female vulva, lateral view. Abbreviations: av = anterior orientated plate; 3iT = 3-combed inner tooth; eT ? external tooth; LT = lateral tooth; mp = molar plate; O = operculum; pL = pectinate lamellae; pv = posterior orientated plate. Scale bars as indicated.

Collum : smooth, laterally not protruding as far as ring 2 (Fig. 4C).

Body rings : ozopores starting at segment 6, marked by a black spot. Located on suture between meso- and metazonite. Rings with smooth, but irregular coriaecous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts elongated, with weak lips, abundant micropunctation especially towards edges (Fig. 4E). Epiproct well-rounded, covering, but not reaching above paraproct (Fig. 4E). Hypoproct inconspicuous (Fig. 4E).

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa. Tarsus with three pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus with a single apical spine and three pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in males.

Female sexual characters. No tarsal pads, antennae shorter than male, only protruding back to ring 2. Female vulva simple, bivalve-like (Fig. 6B). Anterior plate smaller than posterior plate, opening with one row of setae on each plate, close to operculum.

Male sexual characters : tarsal pads present from leg 3 to midbody, small, inconspicuous (Fig. 4D). Coxae 3–7 without coxal processes, but coxae 3–5 swollen (Fig. 3D).

Anterior gonopod sternite massive (Fig. 4F), elongated into a wide, well-rounded triangular lobe (Fig. 4F). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with a large, well-rounded mesal process (FIg. 4F, G). Telopodite with process arising mesally (Fig. 4G), process apically curved with a large triangular projection (Fig. 4G), tip well-rounded, slightly protruding above lateral margin of telopodite (Fig. 4F). Whole telopodite process resembling an even-sided triangle.

Posterior gonopods consisting of three parts, separated by sutures or articulations (Fig. 4H): a basal coxite with a slender coxite projection and a slightly shorter telopodite, efferent duct discharging laterally (FIg. 4H, I). Process of coxite and telopodite standing in same axis (Fig. 4H). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite (Fig. 4H). Basal part of coxite wide, mesally with a large triangular sclerite located on lower level than remaining part (Fig. 4H). Coxite elongated. Efferent duct running at mesal margin of coxite (FIg. 4H, I) before curving to the lateral margin at beginning of telopodite (Fig. 4I). Telopodite as wide as but much shorter than coxite, standing in same axis (FIg. 4H, I), apically membranous, with two slender apical processes both diverging (FIg. 4H, I). Mesal process membranous and wider, lateral process longer, slenderer and sclerotised, efferent duct seems to be ending at base of lateral process (FIg. 4H, I). Base of lateral process with a short, membranous-white projection (Fig. 4I).

The number of segments varies between 47 and 51.

R. mangatsiaka sp. nov. could be found in great numbers in the early morning (7–9 a.m.) on the forest floor of the spiny bush. The otherwise dry spiny bush was still quite wet because of dew. No juveniles were observed. Contrary to other Riotintobolus species, such as R. mandenensis and R. minutus, R. mangatsiaka sp. nov. did not remain stiff like a stick when disturbed, but rolled-up into a spiral (Fig. 4B), a common defence behaviour for juliform millipedes.

1 ♂ holotype, ZFMK MYR9941, Madagascar, South, Lavanono Beach, 25°25.404'S, 044°56.414'E, 27 m, spiny bush at the coast, after rain, leg. Wesener and Schütte, 18.vi.2007.

Paratypes : 16 ♂, 18 ♀, ZFMK MYR942, same data as holotype;

Lavanono, after the type locality, spiny forests directly next to the Lavanono Beach (Fig. 2). Noun in apposition.

Riotintobolus lavanono sp. nov. shares the absence of a projecting epiproct on the telson with R. anomalus, R. antafoky sp. nov., R. bovinus sp. nov., R. tsimelahy sp. nov. and R. mangatsiaka sp. nov., The posterior telopod featuring two slender, sharp projections is only shared with R. bovinus sp. nov., R. mangatsiaka sp. nov. and R. tsimelahy sp. nov. A posterior gonopod separated into three parts is only shared with R. mangatsiaka sp. nov. and R. tsimelahy sp. nov., whose habitus and gonopods look very similar to those of R. lavanono sp. nov. Both species differ in details of the tip of the posterior gonopod and in the colour of their antennae and legs, which are dark grey in R. lavanono sp. nov. and red in both R. mangatsiaka sp. nov. and R. tsimelahy sp. nov. All three species differ by >11% uncorrected p-distance in the COI barcoding gene.

Measurements : male holotype with 47+0 segments, ca. 42 mm long, 4.2 mm wide. Largest females of type series with 46–48+0 segments, up to 58 mm long, 6.4 mm wide.

Colour (in living specimens): Body rings and head grey, appendages black (Fig. 7A). Paraprocts and posterior margins of body segments darker grey to black (Fig. 7B). Ozopore openings highlighted by black spot.

Figure 7. 

Riotintobolus lavanono sp. nov., paratype male (ZFMK MYR942) C anterior gonopod, anterior view; posterior view D telopodite of left posterior gonopod, posterior view E left posterior gonopod, anterior view. Abbreviations: av = paraprocts; Co = collum (ring 1); Cx = coxite; Go = gonopods; Md = basal joints of mandible; Pre = epiproct; St = sternite; sub = hypoproct; T = telopodite; x = lateral swollen process; y = opening of efferent duct; z = apical membranous ‘flag’; numbers refer to leg pair or segment number. Scale bars: 1 mm.

Head : each eye with 28–32 ommatidia in six rows. Incisura lateralis open (Fig. 7A). Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side. Antennae long, protruding back to segment 5. Length of antennomeres: 1<2>3>4=5=6. Terminal antennomere with four large sensory cones located together inside a membranous area. Antennomere 5 and 6 latero-apically with sensilla basiconica.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Endochilarium not dissected.

Mandible : Stipes without projection, well rounded (Fig. 7A). Gnathal lobe not investigated.

Collum : smooth, laterally not protruding as far as ring 2 (Fig. 7A).

Body rings : ozopores starting at segment 6, marked by a black spot. Located on suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts elongated, with weak lips, abundant micropunctation especially towards edges (Fig. 7B). Epiproct well-rounded, covering, but not reaching above paraproct (Fig. 7B). Hypoproct inconspicuous (Fig. 7B).

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa. Tarsus with three pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus with a single apical and four pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in males.

Female sexual characters. No tarsal pads, antennae shorter than male, only protruding back to ring 2. Female vulva simple, bivalve-like.

Male sexual characters : tarsal pads present from leg 3 to midbody, small, inconspicuous. Coxae 3–7 without coxal processes, but coxae 3–5 swollen.

Anterior gonopod sternite massive, elongated into a wide, well-rounded triangular lobe (Fig. 7C). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with a large, well-rounded mesal process. Telopodite with slender process arising mesally (Fig. 7C), process apically curved with a large triangular projection, tip well-rounded, slightly protruding above lateral margin of telopodite.

Posterior gonopods consisting of three parts, separated by sutures or articulations (Fig. 7D): a basal coxite with a slender coxite projection and a shorter telopodite, efferenct duct discharging laterally (Fig. 7E). Process of coxite and telopodite standing in same axis. Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite. Basal part of coxite wide, mesally with a large triangular sclerite located on lower level than remaining part. Coxite elongated. Efferent duct running at mesal margin of coxite before curving to the lateral margin at beginning of telopodite (Fig. 7E). Telopodite half as wide and much shorter than coxite, standing in same axis, apically membranous, with two slender apical processes both diverging (Fig. 7D, E). Mesal process membranous and wider, lateral process bent 90 degrees laterally, longer, slenderer and sclerotised, efferent duct seems to be ending at base of lateral process (Fig. 7E). Base of lateral process with a short, membranous-white projection (Fig. 7E).

Specimens of the same population differing between 45–47 in segment number. Females appear to be more brownish than the more greyish males.

R. lavanono sp. nov. could be found in great numbers after a rainy day in the late afternoon (3–5 p.m.) in a small remnant of spiny bush and under dead Opuntia remains. The specimens were only encountered in an area of a few square meters in view of the coast. Contrary to other Riotintobolus species, such as R. mandenensis and R. minutus, R. lavanono sp. nov. did not remain stiff like a stick when disturbed, but rolled-up into a spiral.

1 ♂ holotype, ZFMK MYR940, Madagascar, Province de Toliara, Makay Mts., forêt de galerie, 21°13'27.5"S, 045°19'35.4"E, 531 m, coll. Jean Noel, 30.xi.2010.

bovinus, after the gonopods which resemble the horns of a cow. Noun in apposition.

Riotintobolus bovinus sp. nov. shares the absence of a projecting epiproct on the telson only with R. anomalus, R. antafoky sp. nov., R. tsimelahy sp. nov., R. mangatsiaka sp. nov. and R. lavanono sp. nov. The posterior telopod featuring two slender, sharp projections is only shared with R. tsimelahy sp. nov., R. mangatsiaka sp. nov. and R. lavanono sp. nov. R. bovinus sp. nov. differs from R. tsimelahy sp. nov., R. mangatsiaka sp. nov. and R. lavanono sp. nov. in a much smaller segment number and size, and strong differences in the posterior telopod, whose telopodite is uniquely shaped with two sharp processes running parallel to one another resembling a bull’s horn. R. bovinus sp. nov. differs by more than 14% uncorrected p-distance in the COI barcoding gene from all other Riotintobolus species.

Measurements : 41+0 segments. Ca. 25 mm long (broken), 2.4 mm wide.

Colour (after 10 years in ethanol): Head and body rings grey, appendages red. Ventral site reddish. Posterior margins of body segments and whole margin of collum black. Anal valves black.

Head : each eye with 24–27 ommatidia in six rows. Incisura lateralis open (Fig. 8A). Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side (Fig. 8B). Antennae short, protruding back to segment 3. Length of antennomeres: 1<2>3>4=5=6, second only slightly longer than 3^rd. Terminal antennomere with four large sensory cones located together inside a membranous area (Fig. 8B). Antennomere 5 and 6 latero-apically with sensilla basiconica.

Figure 8. 

Riotintobolus bovinus sp. nov., male holotype (ZFMK MYR940) A multi-layer photograph, anterior body, lateral view B multi-layer photograph, leg pairs 1–7, ventral view C multi-layer photograph, posterior body end with telson D anterior gonopod, anterior view E anterior gonopod, posterior view F left posterior gonopod, posterior view G left posterior gonopod, anterior view. Abbreviations: Apo = apodeme; av = paraprocts; Co = collum (ring 1); Cx = coxite; Gn = gnathochilarium; Md = basal joints of mandible; Pe = penis; Pre = epiproct; St = sternite; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; numbers refer to leg pair or segment number. Scale bars: 1 mm.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Endochilarium not dissected.

Mandible : Stipes without projection, well rounded (FIg. 8A, B). Gnathal lobe not dissected.

Collum : smooth, laterally not protruding as far as ring 2 (Fig. 8A).

Body rings : ozopores starting at segment 6, marked by a black spot. Located slightly before, but touching suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts without lips, abundant micropunctation especially towards edges (Fig. 8C). Epiproct well-rounded, covering, but not reaching above paraproct (Fig. 8C). Hypoproct inconspicuous (Fig. 8C).

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa and a strongly swollen prefemur. Tarsus with two pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus with a tarsal pad, a single apical and two pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in males.

Female : unknown.

Male sexual characters : tarsal pads absent (Fig. 8B). Coxae 3–7 without coxal processes (Fig. 8B).

Anterior gonopod sternite massive (Fig. 8D), elongated into a wide, well-rounded triangular lobe (Fig. 8D). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with sharp triangular mesal process (Fig. 8D, E). Telopodite with slender process arising mesally (Fig. 8E), process apically curved with a short triangular projection (Fig. 8D), tip well-rounded tip, slightly protruding above lateral margin of telopodite (Fig. 8E).

Posterior gonopods consisting of two parts, separated by an articulation (Fig. 8F): a long coxite and a slightly shorter telopodite, efferent duct discharging apically (Fig. 8F, G). Process of coxite and telopodite standing in same axis (Fig. 8F). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite (Fig. 8F). Basal part of coxite wide, mesally with a large triangular sclerite located on lower level than remaining part (Fig. 8F). Coxite elongated. Efferent duct running at mesal margin of coxite (Fig. 8F, G). Telopodite as wide as but slightly shorter than coxite, standing in same axis (Fig. 8F, G), apically membranous, with two slender apical processes resembling a bull’s horns (Fig. 8F, G). Mesal process wider and longer than lateral process. Efferent duct seems to be ending at base of mesal process (Fig. 8F, G).

Riotintobolus bovinus sp. nov. lives in direct sympatry with another species of the genus, Riotintobolus makayi sp. nov. (Fig. 2).

Holotype : ♂, CASENT 9032794, MGF007, Madagascar, Province Toliara, Antafoky, 80 m, spiny thicket, 23°29'16"S, 44°4'39"E, coll. Frontier project, millipede dig (3 m x 3 m), 14.xi.2001.

Paratypes : 1 ♂, 1 ♀, CASENT 9032794, same data as holotype

Antafoky, after the type locality, spiny forest of Antafoky (Fig. 2). Noun in apposition.

R. antafoky sp. nov. shares the flag-like membranous tip of the posterior gonopod with R. mandenensis, R. minutus, R. aridus, R. anomalus and R. makayi sp. nov. R. antafoky sp. nov. shares the absence of tarsal pads only with R. bovinus sp. nov., and the relatively simple tip of the posterior gonopod only with R. anomalus. R. antafoky sp. nov. differs from the sympatric R. anomalus in details of the posterior gonopod, the absence of a dorsal red stripe, the much longer antenna (protruding back to body ring 5), and the much smaller size (R. anomalus males 45 mm long, 4.3 mm wide, R. antafoky sp. nov. ca. 33 mm long, 3.2 mm wide).

Measurements (holotype): 51+0 segments, ca. 33 mm long (fragmented) and 3.2 mm wide.

Coloration : segments grey, with a dark grey posterior margin, ozopore highlighted by a black spot. Head, antennae and legs dark grey.

Head : each eye with 34 ommatidia in six rows. Incisura lateralis open. Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side. Antennae long, protruding back to segment 5. Terminal antennomere with four large sensory cones located together inside a membranous area. Antennomere 5 and 6 latero-apically with sensilla basiconica.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Endochilarium not dissected.

Mandible : Stipes without projection, well rounded. Gnathal lobe not dissected.

Collum : smooth, laterally not protruding as far as ring 2.

Body rings : ozopores starting at segment 6, marked by a black spot. Located slightly before, but touching suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts without lips, abundant micropunctation especially towards edges. Epiproct well-rounded, covering, but not reaching above paraproct. Hypoproct inconspicuous.

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa and a strongly swollen prefemur. Tarsus with two pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus without a tarsal pad, a single apical and two pairs of ventral spines. Length of midbody legs ca. 0.9 times body diameter in males.

Female : not investigated.

Male sexual characters : tarsal pads absent. Coxae 3–7 without coxal processes, but coxae 6 and 7 flattened, rectangular.

Anterior gonopod sternite massive (Fig. 9A), elongated into a wide, well-rounded triangular lobe (Fig. 9A). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with sharp triangular mesal process (Fig. 9A, B), upper rim bending forward in anterior view (Fig. 9A). Telopodite with strong process arising mesally (Fig. 9B), process apically curved with a short triangular projection (Fig. 9A), tip well-rounded, protruding above lateral margin of telopodite (Fig. 9B).

Figure 9. 

Riotintobolus antafoky sp. nov., holotype (CASENT9032794) A anterior gonopod, anterior view B anterior gonopod, posterior view C telopodite of left posterior gonopod, anterior view D left posterior gonopod, posterior view. Abbreviations: Cx = coxite; St = sternite; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; z = apical membranous ‘flag’. Not to scale.

Posterior gonopods consisting of two parts, separated by an articulation: a long coxite and a slightly shorter telopodite, efferent duct discharging apically (Fig. 9C, D). Process of coxite and telopodite standing in same axis (Fig. 9D). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite (Fig. 9D). Basal part of coxite wide, mesally with a triangular sclerite located on lower level than remaining part (Fig. 9C). Coxite elongated. Efferent duct running at mesal margin of coxite (Fig. 9C, D). Telopodite as wide as but slightly shorter than coxite, standing in same axis (Fig. 9D), apically with a membranous ‘flag’ (Fig. 9C, D). Laterally with sclerotised projection bending laterally and completely surrounded by membranous flag. (Fig. 9C, D). Efferent duct ending at base of process (Fig. 9C, D). Apex laterally with weakly developed finger-shaped process.

this species lives in direct sympatry with the much larger R. anomalus.

1 ♂ holotype, ZFMK MYR7173, Madagascar, Province de Toliara, Makay Mts., forêt de galerie, 21°13'25.1"S, 045°19'36.0"E, 512 m, coll. Jean Noel, 29.xi.2010. Paratypes: 1 ♂, 1 ♀, ZFMK MYR939, Madagascar, Province de Toliara, Makay Mts., forêt de galerie, 21°13'27.5"S, 045°19'35.4"E, 531 m, coll. Jean Noel, 01.xii.2010.

Makayi, after the type locality, the area of Makay (Fig. 2). Noun in apposition.

R. makayi sp. nov. shares the flag-like membranous tip of the posterior gonopod with R. mandenensis, R. minutus, R. aridus, R. anomalus and R. antafoky sp. nov. Riotintobolus makayi sp. nov. shares the wide dorsal stripe, presence of tarsal pads on at least male legs 3–7, and a projecting epiproct only with R. mandenensis, R. minutus, and R. aridus. R. makayi sp. nov. differs from R. mandenensis and R. minutus in the configuration of the flag of the posterior telopod, which consists only of a single fold, while it has two folds in the latter two species. The curved lateral process and the freely ending efferent duct are unique characters for R. makayi sp. nov.

Description (based on male holotype): Measurements: Male holotype: 38+0 segments. Ca. 22 mm long (broken), 2.4 mm wide. Female paratype: 38+0 segments. Ca. 33 mm long (broken), 3.5 mm wide.

Colour (after ten years in ethanol): Head except for light clypeus black, body rings black, appendages red (Fig. 10A, B). Anterior margin of collum light brown. Dorsally with two wide light brown-reddish stripes, divided by a black stripe, all stripes even crossing the epiproct. Anal valves black, margin light brown, hypoproct light brown (Fig. 10C).

Figure 10. 

Riotintobolus makayi sp. nov., male holotype (ZFMK MYR939) A multi-layer photograph, anterior body, lateral view B multi-layer photograph, leg pairs 1–7, ventral view C multi-layer photograph, posterior body end with telson D anterior gonopod, anterior view E anterior gonopod, posterior view F right posterior gonopod, anterior view G right posterior gonopod, posterior view. Abbreviations: Apo = apodeme; av = paraprocts; Co = collum (ring 1); Cx = coxite; Gn = gnathochilarium; Md = basal joints of mandible; Pre = epiproct; St = sternite; T = telopodite; x = lateral swollen process; y = opening of efferenct duct; z = apical membranous ‘flag’; numbers refer to leg pair or segment number. Scale bars: 1 mm.

Head : each eye with 26–28 ommatidia in six rows. Incisura lateralis open (Fig. 10A). Labrum with standard three irregular teeth and a single row of 10–12 stout marginal setae. Clypeus with two setiferous foveolae on each side (Fig. 10B). Antennae short, protruding back to segment 3. Length of antennomeres: 1<2=3=4=5=6. Terminal antennomere with four large sensory cones located together inside a membranous area (Fig. 10B). Antennomere 5 and 6 latero-apically with sensilla basiconica.

Gnathochilarium : lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Endochilarium not dissected.

Mandible : Stipes without projection, well rounded (Fig. 10A, B). Gnathal lobe not dissected.

Collum : smooth, laterally not protruding as far as ring 2 (Fig. 10A).

Body rings : ozopores starting at segment 6, located slightly before, but touching suture between meso- and metazonite. Rings with smooth, but irregular coriaceous surface, ventrally on metazona with transverse ridges.

Telson : paraprocts with lips, abundant micropunctation especially towards edges (Fig. 10C). Epiproct well-rounded, reaching slightly above paraproct with a massive process (Fig. 10C). Hypoproct inconspicuous (Fig. 10C).

Legs : leg 1 with a large cylindrical coxa, twice as long as other podomeres. Tarsus with three pairs of ventral spines and an apical spine beyond claw. Leg 2 with an elongated coxa. Tarsus with two pairs of ventral spines and a short apical spine. Midbody legs with a rectangular coxa, as long as other podomeres. Each podomere ventrally with a single or a pair of apical setae, tarsus of leg 3– midbody with a tarsal pad, a single apical and three or four pairs of ventral spines (Fig. 10B). Legs on posterior part of body without a tarsal pad and two or three pairs of ventral spines. Length of midbody legs ca. 1.2 times body diameter in male.

Female sexual characters. No tarsal pads, antennae even shorter than male, only protruding back to collum. Female vulva simple, bivalve-like.

Male sexual characters : tarsal pads present from leg 3 to midbody (Fig. 10B). Coxae 3–7 without coxal processes (Fig. 10B).

Anterior gonopod sternite massive, elongated into a wide, well-rounded, triangular lobe (Fig. 10D). Sternite in anterior view well-visible, without discernible apodemes, protruding almost as high as coxal processes. Coxite with sharp triangular mesal process (Fig. 10D, E). Telopodite with slender process arising mesally (Fig. 10E), process apically curved with a strong triangular projection (Fig. 10D), tip well-rounded tip, protruding above lateral margin of telopodite (Fig. 10C).

Posterior gonopods consisting of two parts, separated by an articulation: a long coxite and a slightly shorter telopodite, efferent duct discharging apically (Fig. 10G). Process of coxite and telopodite standing in same axis (Fig. 10F, G). Pair of posterior gonopods located parallel to each other, connected by a small, sclerotised and visible sternite (Fig. 10G). Basal part of coxite wide, mesally with a triangular sclerite located on lower level than remaining part (Fig. 10G). Coxite elongated. Efferent duct running at mesal margin of coxite (Fig. 10F, G). Telopodite as wide as but slightly shorter than coxite, standing in same axis, apically with a membranous ‘flag’ (Fig. 10F, G). Laterally with sclerotised projection with a slender mesad-orientated branch (Fig. 10F, G). Membranous ‘flag’ longer than lateral process. Efferent duct ending freely, projecting above ‘flag apically (Fig. 10F, G).

this species lives in direct sympatry with Riotintobolus bovinus sp. nov. The female carried eggs in the body.