Riotintobolus lavanono,

Wesener, Thomas, 2020, Ecotone shifts in southern Madagascar: first barcoding data and six new species of the endemic millipede genus Riotintobolus (Spirobolida, Pachybolidae), ZooKeys 953, pp. 1-29: 1

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Riotintobolus lavanono

sp. nov.

Riotintobolus lavanono  sp. nov. Figure 7View Figure 7

Material examined.

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,

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


Lavanono, after the type locality, spiny forests directly next to the Lavanono Beach (Fig. 2View Figure 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. 7AView Figure 7). Paraprocts and posterior margins of body segments darker grey to black (Fig. 7BView Figure 7). Ozopore openings highlighted by black spot.

Head: each eye with 28-32 ommatidia in six rows. Incisura lateralis open (Fig. 7AView Figure 7). 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. 7AView Figure 7). Gnathal lobe not investigated.

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

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. 7BView Figure 7). Epiproct well-rounded, covering, but not reaching above paraproct (Fig. 7BView Figure 7). Hypoproct inconspicuous (Fig. 7BView Figure 7).

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. 7CView Figure 7). 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. 7CView Figure 7), 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. 7DView Figure 7): a basal coxite with a slender coxite projection and a shorter telopodite, efferenct duct discharging laterally (Fig. 7EView Figure 7). 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. 7EView Figure 7). Telopodite half as wide and much shorter than coxite, standing in same axis, apically membranous, with two slender apical processes both diverging (Fig. 7D, EView Figure 7). 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. 7EView Figure 7). Base of lateral process with a short, membranous-white projection (Fig. 7EView Figure 7).

Intraspecific variation.

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

Live observations.

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.