Attemsostreptus reflexus Akkari & Enghoff, 2019

Enghoff, Henrik, Ngute, Alain S. K., Kwezaura, Revocatus L., Laizzer, Richard L., Lyatuu, Herman M., Mhagawale, Waziri, Mnendendo, Hamidu R. & Marshall, Andrew R., 2024, A mountain of millipedes XI. The trachystreptoform spirostreptids of the Udzungwa Mountains, Tanzania (Diplopoda, Spirostreptida, Spirostreptidae), European Journal of Taxonomy 918 (1), pp. 1-50 : 6-13

Attemsostreptus reflexus Akkari & Enghoff, 2019

Figs 2 , 3A–B , 4–7

Attemsostreptus reflexus Akkari & Enghoff, 2019: 3

Diagnosis

Easily distinguishable from congeners by the extremely long lateroapical metaplical process bent in a U-shape ( Figs 3B , 7A ).

Material examined (total 25 ♂♂, 33 ♀♀, 78 juvs)

TANZANIA – Morogoro Region, Kilombero District, Udzungwa Mountains National Park , Mwanihana • 4 ♂♂, 9 ♀♀, 11 juvs; Njokamoni ; 07°50′35.9″ S, 36°52′38.6″ E; 474 m a.s.l.; 20 Dec. 2021; A. Ngute, R. Malanda, W. Mhagawale, A. Marshall leg.; FoRCE Plot 10, closed canopy; COLL. NHMD - ACC.NO. 2022-EN-003; NHMD 621851 • 1 ♂, 12 juvs; same collection data as for preceding; 07°50′31.9″ S, 36°52′471″ E; 401 m a.s.l.; 21 Dec. 2021; FoRCE Plot 11, open canopy; NHMD 621852 • 1 ♂, 4 ♀♀, 1 juv.; same collection data as for preceding; 07°50′33.2″ S, 36°52′54.9″ E; 357 m a.s.l.; 22 Dec. 2021; FoRCE Plot 12, open canopy; NHMD 621853 • 1 ♀, 12 juvs; Sonjo ; 07°48′22.6″ S, 36°52′17.0″ E; 579 m a.s.l.; 14 Dec. 2021; A. Ngute, R. Malanda, H. Mnendendo, W. Mhagawale, A. Mpoto, A. Marshall leg.; FoRCE Plot 13, closed canopy; NHMD 621854 • 2 juvs; same collection data as for preceding; 07°48′24.4″ S, 36°52′21.7″ E; 589 m a.s.l.; 15 Dec. 2021; FoRCE Plot 14, open canopy; NHMD 621855 • 1 juv.; same collection data as for preceding; 07°48′22.3″ S, 36°52′32.2″ E; 595 m a.s.l.; 17 Dec. 2021; FoRCE Plot 15, open canopy; NHMD 621856 • 3 ♂♂, 2 ♀♀, 1 juv.; Sanje ; 07°46′43.4″ S, 36°54′04.3″ E; 504 m a.s.l.; 26 Feb. 2022; A. Ngute, E. Kivambe, R. Malanda, H. Mnendendo, W. Mhagawale, M. Mpoto, A. Marshall leg.; FoRCE Plot 24, open canopy; NHMD 621857 • 3 ♂♂; same collection data as for preceding; 07°46′47.6″ S, 36°54′07.6″ E; 495 m a.s.l.; 23 Feb. 2022; FoRCE Plot 25, open canopy; NHMD 621858 • 1 ♀, 2 juvs; same collection data as for preceding; 07°46′52.1″ S, 36°54′11.8″ E; 421 m a.s.l.; 20 Feb. 2022; A. Ngute, E. Kivambe, R. Malanda, W. Mhagawale, H. Mnendendo, A. Marshall leg.; FoRCE Plot 92, open canopy; NHMD 621859 • 1 ♀; same collection data as for preceding; 07°47′03.0″ S, 36°54′05.8″ E; 410 m a.s.l.; A. Ngute, E. Kivambe, R. Malanda, W. Mhagawale, H. Mnendendo, A. Marshall leg.; FoRCE Plot 93, open canopy; NHMD 621860 . – Morogoro Region, Kilombero District, Udzungwa Mountains National Park • 1 ♂, 2 juvs; 07°49′21.7″ S, 36°58′57.1″ E; 283 m a.s.l.; 30 Jan. 2020; A. Ngute and A.R. Marshall leg.; micro-FoRCE plot 12, closed forest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 621861 • 2 ♀♀, 1 juv.; same collection data as for preceding; 07°49′13.2″ S, 36°58′57.1″ E; 334 m a.s.l.; 25 Feb. 2020; micro-FoRCE plot 17; open forest; NHMD 621862 • 2 ♂♂, 4 juvs; same collection data as for preceding; 07°50′14.5″ S, 36°53′07.8″ E; 358 m a.s.l.; 13 Feb. 2020; micro-FoRCE plot 18; closed forest; NHMD 621863 • 2 ♀♀; same collection data as for preceding; 07°49′46.5″ S, 36°53′27.9″ E; 347 m a.s.l.; 27 Feb. 2020; micro-FoRCE plot 20, open forest; NHMD 621864 • 1 juv.; same collection data as for preceding; 07°48′53.2″ S, 36°54′13.9″ E; 282 m a.s.l.; 5 Mar. 2020; micro-FoRCE plot 22, open forest; NHMD 621865 • 2 ♂♂, 3 ♀♀; same collection data as for preceding; 07°49′03.6″ S, 36°57′33.9″ E; 280 m a.s.l.; 16 Mar. 2020; micro-FoRCE plot 27, closed forest; NHMD 621866 • 1 ♀, 7 juvs; same collection data as for preceding; 07°47′43.1″ S, 36°53′57.6″ E; 331 m a.s.l.; 19 Feb. 2020; micro-FoRCE plot 28, open forest; NHMD 621867 • 1 ♂, 4 ♀, 9 juvs; same collection data as for preceding; 07°47′01.1″ S, 36°54′13.9″ E; 381 m a.s.l.; 15 Feb. 2020; micro-FoRCE plot 30, closed florest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 621868 . – Morogoro Region, Kilombero District, Magombera Nature Reserve • 1 ♂, 3 ♀, 10 juvs; 07°48′53.0″ S, 36°58′29.0″ E; 285 m a.s.l; 5 Mar. 2018; A.R. Marshall leg.; half-FoRCE plot 2, closed forest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 621869 • 1 ♂; same collection data as for preceding; 07°48′59.4″ S, 36°57′55.6″ E; 280 m a.s.l.; 14 Mar. 2018; half-FoRCE plot 3, closed forest; NHMD 621870 • 1 ♂, 1 ♀; same collection data as for preceding; 07°49′03.0″ S, 36°57′35.2″ E; 284 m a.s.l.; 15 Mar. 2018; half-FoRCE plot 4, closed forest; NHMD 621871 • 1 ♂, 1 ♀, 1 juv.; same collection data as for preceding; 07°48′30.9″ S, 36°59′33.1″ E; 287 m; 19 Feb. 2018; half-FoRCE plot 37, open forest; NHMD 621872 • 1 juv.; same collection data as for preceding; 07°48′21.5″ S, 36°59′47.5″ E; 287 m a.s.l.; 17 Feb. 2018; half-FoRCE plot 38, open forest; NHMD 621873 • 2 ♂♂; same collection data as for preceding; 07°48′53.0″ S, 36°58′29.0″ E; 285 m; 27 Jun. 2019; A. Ngute and A.R. Marshall leg.; half-FoRCE plot 2, closed forest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 621874 • 1 ♂, 6 ♀♀, 1 juv.; same collection data as for preceding; 07°49′03.0″ S, 36°57′36.2″ E; 284 m a.s.l.; half-FoRCE plot 4, closed forest; NHMD 621875 .

Descriptive notes (males)

This species was recently described and illustrated with photographs ( Akkari & Enghoff 2019). We here provide descriptive notes based on the new material from the Udzungwa Mountains.

SIZE. Males: vertical diameter 3.6–4.6 mm; 52–54 podous rings, no apodous rings in front of telson.

COLLUM ( Fig. 4A ). With smooth surface, with 3 strong lateral striae/carinae, above and below these a few indistinct ones, a short stria/carina in each of the spaces between the main ones.

BODY RINGS ( Fig. 5 ). Anterior part of prozonite (pz) with fine, dense transverse furrows; central part with an irregular mesh of cells delimited by fine ridges; posterior row of cells longer and more regular; surface of cells pitted; suture (sut) between pro- and metazonite delimited by sharp edge, a row of tiny, sub-quadrate cells between edge and posterior row of prozonital cells ( Fig. 5E ). Metazonite (mz) with densely set sharp ridges throughout perimeter, anterior part of ridges with string-of-bead-like sculpture, surface between ridges pitted. Ozopore ( Fig. 5D, F ) ca ⅓ metazonite length behind suture, oval, surrounded by narrow concentric cytoscutes and with a conical sclerotized plug. No sigilla.

TELSON ( Fig. 4B–D ). Preanal ring (pr) with coarsely rugose surface. Anal valves (av) with strongly developed submarginal lips; apical and mesal surface of lips smooth, outer surface of lips and rest of valves with coarsely rugose microsculpture.

LEGS ( Fig. 6 ). Male legs with well-developed ventral pads on postfemur and tibia ( Fig. 6E ). First pair ( Fig. 6A–D ): coxosternum with lateral groups of a few setae (cxs). Prefemoral lobes (pfl) in ventral view equilaterally triangular, pointed, covered with minute short spines ( Fig. 6D ).

GONOPODS ( Figs 3B , 7 ). As described by Akkari & Enghoff (2019). Notable details include 1) coxal metaplica (mp) apically folded back on anterior side, forming a “roof” or canopy (mr) over gonocoel; 2) distomesal metaplical spine (mcs) long, protruding at angles (ca 40°) from mesal metaplical margin; 3) tip of telopodite bifurcate, one branch (sl) lamellar, with transverse, slightly serrated tip, the other (solenomere, slm) acuminate.

Descriptive notes (females)

Diameter up to 5.1 mm; 52–55 podous rings, no apodous rings in front of telson.

Distribution and habitat

In the Udzungwa Mountains, it was found on 25 plots of the FoRCE experiment of which 11 are with open canopy and 14 with closed canopy. The total altitudinal range is 280–595 m a.s.l. Although abundant in the FoRCE material, the species is absent from older collections, maybe an indication that it is a recent immigrant to the Udzungwa Mountains. The only other precise record of the species is from Kimboza Forest Reserve in the Uluguru Mountains, Tanzania ( Akkari & Enghoff 2019).

Figures

Fig. 2. Map of the Udzungwa Mts showing collection areas for trachystreptoform Spirostreptidae Brandt, 1833. Yellow symbols: Attemsostreptus spp.; dots: A. reflexus, diamond: A. cataractae Enghoff sp. nov., cross: A. leptoptilos Enghoff sp. nov., triangles; A. julostriatus Enghoff sp. nov. Red symbols: Lophostreptus spp.; dots:L. tersus, crosses:L. magombera Enghoff sp. nov. Blue dots:Udzungwastreptus marianae Enghoff gen. et sp. nov. Based on Marshall et al. (2010: fig. 1).

Fig. 3. Trachystreptoforms from the Udzungwa Mts.A. Attemsostreptus reflexus Akkari & Enghoff, 2019, ♂ (Mwanihana, Njokamoni, NHMD 621851). B. Same, close-up of gonopod in situ. C. Attemsotreptus julostriatus Enghoff sp. nov., holotype, ♂ (NHMD 621878). D. Attemsostreptus leptoptilos Enghoff sp. nov., paratype, ♂ (NHMD 621877). E–F. Lophostreptus tersus (Cook, 1896). E. Two living females from Magombera Nature Reserve (NHMD 1184642). F. Male from Magombera Nature Reserve (NHMD 1184640). G. Lophostreptus magombera Enghoff sp. nov., paratype, ♂ (NHMD 621886). H. Udzungwastreptus marianae gen. et sp. nov., paratype, ♂ (NHMD 621893). Scale bars: A, C–D, F–H = 2 mm; B = 0.5 mm; E not to scale. Photos: S.G. Selvantharan (A–D, F–H), A.R. Marshall (E).

Fig. 4. Attemsostreptus reflexus Akkari & Enghoff, 2019, male from Udzungwa Mts (Mwanihana, Njokamoni, NHMD 621851). A. Head, collum and rings 2–6, lateral view. B–D. Telson and ultimate ring. B. Lateral view. C. Posterior view, incl. legs of ultimate ring. D. Dorsal view. Abbreviations: av = anal valve; pr = preanal ring. Scale bars = 0.5 mm.

Fig. 5. Attemsostreptus reflexus Akkari & Enghoff, 2019, male from Udzungwa Mts (Mwanihana, Njokamoni, NHMD 621851), midbody ring. A. In toto. B. Prozonite, lateral part. C. Prozonital sculpture just in front of suture. D. Metazonital sculpture just behind suture. E. Same, close-up. F. Ozopore. Abbreviations: mz = metazonite; oz = ozopore; pz = prozonite; sut = suture. Scale bars: A = 0.2 mm; B = 0.1 mm; C, E = 0.02 mm; D = 0.05 mm; F = 0.005 mm.

Fig. 6. Attemsostreptus reflexus Akkari & Enghoff, 2019, male from Udzungwa Mts (Mwanihana, Njokamoni, NHMD 621851). A–D. First pair of legs. A. Posterior view. B. Ventral view. C. Sublateral view. D. Peglike setae/sensilla from prefemur.E. Midbody leg, arrows pointing at soft pads on postfemur and tibia. Abbreviations: cxs = coxosternal setae; pfl = prefemoral lobe. Scale bars: A–B, E = 0.2 mm; C = 0.1 mm; D = 0.02 mm.

Fig. 7. Attemsostreptus reflexus Akkari & Enghoff, 2019, male from Udzungwa Mts (Mwanihana, Njokamoni, NHMD 621851). A. Gonopods except right telopodite, anterior view; inset: close-up of mesal coxal spines. B. Tip of left telopodite. C–D. Tip of right telopodite. E–G. Right telopodite. E. Sublateral view.F. anterior view; striped rectangle: area obscured by mounting tape.G. Posterior view. Abbreviations: atp = antetorsal process (highlighted in yellow in F); ba = basomere; lap = lateroapical metaplical process; map = mesapical metaplical process; mcs = mesal coxal spine; mp = metaplica; mr = metaplical “roof”; pp = proplica; px = paracoxite; sl = serrate lamella; slm = solenomere. Scale bars: A, E–G = 0.2 mm; B–D = 0.05 mm.