Lophostreptus tersus ( Cook, 1896 )

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 : 24-34

Lophostreptus tersus ( Cook, 1896)

Figs 1– 2 , 3E–F , 14–17

Ptilostreptus tersus Cook, 1896: 57 .

Lophostreptus ptilostreptoides Carl, 1909: 321 , syn. nov. (tentatively suggested by Carl 1909: 317).

Lophostreptus regularis Attems, 1909: 31 , synonymized with L. ptilostreptoides by Krabbe (1982: 258).

Lophostreptus malleolus Kraus, 1958: 12 , synonymized with L ptilostreptoides by Demange & Mauriès (1975: 79).

Lophostreptus tersus – Attems 1914: 143.

Lophogonus ptilostreptoides – Demange & Mauriès 1975: 78.

Diagnosis

Differs from congeners and all other trachystreptoform spirostreptids by the complex and highly characterictic lateroapical metaplical process (lap): sharply bent laterad, in the shape of a long, twisted slipper with a rounded mesal ‘heel’. Very similar to that of the sympatric L. magombera sp. nov., but larger ( Fig. 17 ).

Material examined

Material from Udzungwa Mountains (total 16 ♂♂, 27 ♀♀, 24 juvs)

TANZANIA – Morogoro Region, Kilombero District, Magombera Nature Reserve • 2 ♂♂, 4 ♀♀; 07°49′03.6″ S, 36°58′40.0″ E; 279 m a.s.l.; 22 Jan. 2020; A. Ngute and A.R. Marshall leg.; micro-FoRCE plot 10, open forest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 1184638 • 1 ♀, 1 juv.; same collection data as for preceding; 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; NHMD 1184639 • 3 ♂♂, 5 ♀♀, 2 juvs; same collection data as for preceding; 07°49′28.0″ S, 36°58′55.2″ E; 271 m a.s.l.; 31 Jan. 2020; micro-FoRCE plot 13, open forest; NHMD 1184640 • 3 ♂♂; Udzungwa Mountains National Park ; 07°50′56.4″ S, 36°53′00.4″ E; 325 m a.s.l.; 2 Mar. 2020; A. Ngute and A.R. Marshall leg.; micro-FoRCE plot 26, open forest; NHMD 1184641 • 2 ♀♀, 1 juv.; same collection data as for preceding; 07°48′30.9″ S, 36°50′33.1″ E; 287 m a.s.l.; 14 Jul. 2019; A. Ngute and A.R. Marshall leg.; half-FoRCE plot 37, open forest; COLL. NHMD. ACC.NO. 2020-EN-002; NHMD 1184642 • 1 juv. ♂; same collection data as for preceding; Ngulumilo ; 07°49′13.2″ S, 36°59′10.3″ E; 285 m a.s.l., 23 Aug. 2021; A. Ngute, E. Kivambe, R. Malanda, W. Mhagawale, H. Mnendendo, A. Marshall leg.; FoRCE plot 1, closed canopy; COLL. NHMD - ACC.NO. 2022-EN-003; NHMD 1184643 . – Morogoro Region, Kilombero District, Udzungwa Mountains National Park, Mwanihana • 1 ♂, 5 ♀♀, 7 juvs; Njokamoni ; 07°50′35.8″ S, 36°52′38.5″ 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 1184644 • 2 ♀♀, 2 juvs; same collection data as for preceding; Njokamoni ; 07°50′31.9″ S, 36°52′47.1″ E; 401 m a.s.l.; 21 Dec. 2021; FoRCE plot 11, open canopy; NHMD 1184645 • 1 ♀; same collection data as for preceding; Njokamoni ; 07°50′33.1″ S, 36°52′54.9″ E; 357 m a.s.l.; 22 Dec. 2021; FoRCE plot 12, open canopy; NHMD 1184646 • 1 ♂, 1 ♀, 1 juv.; same collection data as for preceding; Sonjo ; 07°48′24.4″ S, 36°52′21.7″ E; 589 m a.s.l.; 15 Dec. 2021; A. Ngute, R. Malanda, H. Mnendendo, W. Mhagawale, A. Mpoto, A. Marshall leg.; FoRCE plot 14, open canopy; NHMD 1184647 • 1 ♂, 2 ♀♀, 7 juvs; same collection data as for preceding; Sonjo ; 07°48′22.3″ S, 36°52′32.2″ E; 595 m a.s.l.; 17 Dec. 2021; FoRCE plot 15, open canopy; NHMD 1184648 • 1 ♀, 1 juv.; same collection data as for preceding; Sonjo ; 07°48′22.3″ S, 36°52′32.2″ E; 541 m a.s.l.; 18 Aug. 2021; A. Ngute, E. Kivambe, W. Mhagawale, H. Mnendendo, A. Mwakisoma, A. Marshall leg.; FoRCE plot 95, open canopy; NHMD 1184649 • 1 ♂, 1 ♀; same collection data as for preceding; Mizimu ; 07°48′34.0″ S, 36°51′17.0″ E; 900 m a.s.l.; 12 Dec. 2021; FoRCE plot 17, open canopy, subplot 1; NHMD 1184650 • 2 ♂♂, 1 ♀; same collection data as for preceding; Sanje ; 07°46′47.6″ S, 36°54′07.6″ E; 495 m a.s.l.; 23 Feb. 2022; A. Ngute, E. Kivambe, R. Malanda, H. Mnendendo, W. Mhagawale, M. Mpoto, A. Marshall leg.; FoRCE plot 25, open canopy; NHMD 1184651 • 1 ♂; same collection data as for preceding; Sanje ; 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 1184652 • 1 ♀, 1 juv.; same collection data as for preceding; Sanje ; 07°47′03.0″ S, 36°54′05.8″ E; 410 m a.s.l.; 17 Feb. 2022; A. Ngute, E. Kivambe, W. Mhagawale, H. Mnendendo, M. Mpoto, A. Marshall leg.; FoRCE plot 93, open canopy; NHMD 1184653 . – Morogoro Region, Ecological Monitoring Centre Mang’ula • 1 ♂; 07°50′44.9″ S, 36°53′28.2″ E; 339 m a.s.l.; 20 Mar. 2013; T. Pape and N. Scharff leg.; hand-collected; NHMD 1184654 .

Material from other places than Udzungwa Mountains (total 11 ♂♂, 30 ♀♀, 4 juvs)

TANZANIA • 3 ♂♂, 14 ♀♀; Arusha, Momella ; 1300 m a.s.l.; ult. Dec. 1975; L. and T. Nielsen leg.; NHMD 1184655 • 1 ♂; ca 10 km E of Arusha; 1300 m a.s.l.; 19 Jan. 1971; H. Enghoff, O. Lomholdt and O. Martin leg.; NHMD 1184656 • 4 ♂♂; Morogoro Region, Morogoro District, Kimboza Forest Reserve ; 07°01′ S, 37°48′ E; Jan.–Mar. 1994; Frontier Tanzania leg.; NHMD 1184657 • 1 ♂; Morogoro Region, Morogoro District, Mindu Mountains ; 06°29′ S, 37°21′ E; 1200 m a.s.l.; 24 Dec. 1983; J. Kielland leg.; VMNH 112023 • 1 ♂, topotype of Ptiostreptus tersus ; Dodoma Region, Mpwapwa District, Wota Forest Reserve ; Apr. 1984; J. Kielland leg.; VMNH 112024 • 1 ♂, 6 ♀♀, syntypes of Lophostreptus regularis ; Kilimandjaro, Kibonoto, Stepp-Kulturzon ; 1000–1900 m a.s.l.; Oct. 1905; Y. Sjöstedt leg.; also 1 ♂ of L. neglectus ; NHRS-TOBI 000005480 • 4 ♀♀, syntypes of L. regularis ; Kilimandjaro, Kibonoto, Massaistäppen ; 1000 m a.s.l.; 23 Aug. 19905; Y. Sjöstedt leg.; NHRS-TOBI 000005478 • 1 ♀, syntype of L. regularis ; Usambara , Tanga; Jun. 1905; Y. Sjöstedt leg.; NHRS-TOBI-000005482 • 4 ♀♀, syntypes of L. regularis ; Kilimandjaro, Kibonoto ; Nov. 1905; Y. Sjöstedt leg.; “under multnande blad i bananfarmerkulturzon” [under decaying leaves in banana farm cultural zone]; NHRS-TOBI-000005476 • 4 ♀♀, syntypes of L. regularis ; Kilimandjaro, Kibonoto ; 1300 m a.s.l.; 1905; Y. Sjöstedt leg.; “I förnan under nedfallna plantanblad” [in förna under fallen plantain leaves]; NHRS-TOBI-000005479 • 1 ♀, syntype of L. regularis ; Kilimandjaro, Kibonoto ; 1905; Y. Sjöstedt leg.; Mischwald–Kulturzone; NHRS-TOBI-000005481 • 4 ♀♀, 4 anamorphic juvs, syntypes of L. regularis ; Kilimandjaro, Kibonoto ; Nov. 1905; Y. Sjöstedt leg.; Kulturzon; NHRS-TOBI-0000077 .

Descriptive notes on males from Udzungwa Mountains

SIZE. Length 52–58 mm; vertical diameter 3.8–4.5 mm; adults and epimorphic juveniles with 48–52 podous rings ( Figs 14 , 18 ), no apodous rings in front of telson.

COLOUR. Live colour ( Fig. 3E ) overall grey, legs orange yellow.After 2–3 years in alcohol head, antennae, collum, body rings 2–7, telson and dorsal part of remaining body rings blackish brown (head capsule sometimes broadly yellow at labral margin); lateral and ventral part of remaining body rings from well above ozopore level rarely same colour, usually rusty red-brown with contrasting small black spots covering ozopores; posterior edge dark amber; legs medium brown.

HEAD ( Fig. 15A–B ). Almost smooth below antennae, longitudinally wrinkled near anterior edge, finely punctate between antenna and eyes. Vertex very densely and rather coarsely punctuate, with a clearly demarcated parietal furrow. Eyes not reaching mesal of antennal socket, ca 40 ommatidia in ca 6 horizontal and ca 12 vertical rows. Antennae reaching posterior margin of 2 nd –3 rd body ring. Antennomeres 3–5 strongly narrowed at base.

COLLUM ( Fig. 15A–B ). Not modified for accommodation of antennae, densely punctuate, finely dorsally, more coarsely towards the sides; along posterior margin a row of quite short, weak furrows and carinae which towards the sides gradually reach further forwards. Lateral lobes much narrower than dorsal part, not expanded, traversed by 3 or 4 anteriorly strongly ascending carinae/furrows of which uppermost is strongest and almost straight, reaching anterior margin above eye level, anterior corner rectangular, posterior corner more rounded, margins straight.

BODY RINGS ( Fig. 15A, C–D ). Prozonites (pz) in anterior part (ca half) with very fine ring furrows which further back give place to an irregular cell structure; posterior part (ca 20%) especially dorsally delimited by clear line, with a more regular pattern of larger cells. Suture between pro- and metazonites straight, simple. Metazonites (mz) with clear constriction a little behind suture, with numerous simple keels which at least dorsally reach from suture, across constriction and until posterior ring margin; ca 25 keels between dorsal midline and ozopore in a male of 4.3 mm diameter; in larger specimens keels on lateral flanks of anterior segments projecting as short spines beyond posterior metazonital margin (as described by Attems 1914). Ozopores (oz) small, a little before middle of metazonite. A row of large sigilla. Sternites transversely striate.

TELSON ( Fig. 15E–G ). Preanal ring (pr) regularly and densely grainy-rugose. Anal valves (av) overall with same sculpture, strongly vaulted, their mesal margins slightly raised as low rims, smooth, meeting in midline, paralleled more laterally by much higher lips with smooth edge; distance between lips and mesal margin equal to or larger than height of lips; area between mesal margin and keel with weaker sculpture than main part of valve.

LEGS. Short, length ca 0.6×body diameter. No ventral pads. First pair ( Fig. 16 ): coxosternum with lateral groups of a few long setae (cxs) and more mesally with large groups of numerous long setae (cxs) next to prefemoral lobes. Prefemoral lobes (pfl) rounded-rectangular, with a few apicomesal setae (aps), otherwise naked.

GONOPODS ( Fig. 17 ). As described by Cook (1896), Carl (1909), Attems (1909), Kraus (1958) and Krabbe (1982). Notable features include:

– the very distinctive lateroapical metaplical process (lap); shaped like an irregular, twisted slipper, somewhat constricted ca at midlength

– the rounded mesapical metaplical process (map); forming the “heel of the slipper”

– a distinct “knee” (kn) on the free part of the telopodite, ca ⅓ from its emergence from the gonocoel

– the very large, palette-shaped appendage (pal) at the transition between the middle and the distal ⅓ of the telopodite

– the four-pronged tip of the telopodite, the solenomere (slm) being flanked by one tongue-shaped (tp1) and two slender processes (tp2, tp3)

Descriptive notes on females from Udzungwa Mountains

Vertical diameter up to 5.3 mm. Adults and epimorphic juveniles with 48–52 podous rings ( Figs 14 , 18 ), no apodous rings in front of telson.

Vulvae quite well sclerotized; operculum small, tounge-shaped; valves transversely wrinkled, meeting in very oblique line, mesal valve much larger than lateral valve, resemble the vulva ascribed to “ Lophostreptus? regularis ” by Brölemann (1920: fig. 30).

Distribution and habitat

Quite widespread in the northern half of Tanzania and in southern Kenya ( Enghoff et al. 2016, as L. ptilostreptoides ). In the Udzungwa Mountains, it was found on 16 plots of the FoRCE experiment of which 13 are with open canopy and only three with closed canopy. The total altitudinal range is 271–900 m a.s.l., but 15 of the 16 plots are below 600 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. Some earlier reports of the species suggest that it is at least in part synanthropic: banana plantation ( Carl 1909, as L. ptilostreptoides ) cultural zone ( Attems 1909, as L. regularis ), botanical garden ( Kraus 1958, as L. malleolus ), around houses ( Mwabvu & VandenSpiegel 2009, as L. ptilostreptoides ).

Notes on a topotype of Ptilostreptus tersus Cook, 1896

The male from Dodoma Region, Mpwapwa District, Wota Forest Reserve (VMNH 112024) can be regarded as a topotype, at least a near-topotype of Ptilostreptus tersus – type locality “Mpapua, German East Africa ” according to Cook (1896).

Compared side-by-side to a male from the Udzungwa Mountains, the P. tersus topotype differs in a few gonopod details:

– the coxa on the whole appears relatively slenderer

– the mesapical metaplical process (map) forms a small hook at its meso-basal corner

– the part of the telopodite from the torsus until the large palette-like appendage (pal) is straighter

– the palette-like appendage (pal) is narrower, more than twice as long as broad

– the part of the telopodite distal to the palette-like appendage (pal) appears longer

See further under “Remarks” below.

Notes on type material of Lophostreptus regularis Attems, 1909

Syntypical material of this nominal species belonging to NRMS (see Material examined) was studied, and Nesrine Akkari kindly provided detailed information on the syntypes belonging to NHMW, including photos of the specimens and of Attems’ microscope slides. Further syntypes are present in Museum für Naturkunde, Berlin ( Moritz & Fischer 1974: 371), but have not been examined.

The syntypes from NRMS include two adult males. One of them lacks its posterior part and has a body diameter of 3.0 mm. Notably, the legs have no traces of ventral pads (contra Attems 1909: 32, fig. 57). One of the males has gonopods exactly like those of L. tersus , but the second belongs to L. neglectus . This species is also “trachystreptoform” but is much smaller than Lophostreptus tersus (= L. regularis ): 46 podous rings, no apodous rings, vertical diameter 2.4 mm. Also this species lacks ventral pads on the legs.

The syntypes from NHMW include two undissected males which, according to the photos provided by Nesrine Akkari, belong to L. regularis (the characteristic process lap is clearly visible on the photos). On the other hand, Attems’ original slides are rather confusing: slide “NHMW MY 8871 (ex 4076)”: contains a set of gonopods which clearly belong to L. tersus . However, among the slides labelled “NHMW MY 4076”, one contains a set of typical L. tersus gonopods (in very poor condition), two contain legs with pads on postfemur and tibia (one of the legs corresponds in all details to Attems’ fig. 57), and one contains a full set of gonopods which obviously belong to L. neglectus . Considering that this species, like all examined males of L. tersus , lacks postfemoral and tibial pads, the origin of the pad-bearing legs in the slide from the NHMW MY 4076 series remains a mystery. In summary, it seems that the NHMW syntypes of L. regularis represent three species: an unknown species with padded legs. L. tersus , and a probably undescribed “trachystreptoform”.

In couplet 5 of the key to species of Lophostreptus by Attems (1938), L. regularis is separated from L. tersus and L. ptilostreptoides by having a “femoral” spine on the gonopods, whereas the two latter species have no such spine. This must be a mistake – nothing like a femoral spine is mentioned or illustrated in the original description of L. regularis .

Remarks

In Table 2 , the dimensions of L. tersus according to previous authors are shown, together with data on examined specimens from various localities.

Mwabvu & VandenSpiegel (2009) recorded L. ptilostreptoides from the Taita Hills in S Kenya and presented a drawing of the gonopods, but they gave no further descriptive details.

The colour has been described quite differently, but given the known, variable effect of preservation, already evident after a few years, cf. the description of Udzungwa material, such differences are not regarded significant.

The overall variation in ring number (45–52) is largely covered by Udzungwa specimens (48–52), and this provides no basis for subdividing the material into several species. In terms of body diameter, the variation is considerable (2.5–4.5 mm for adult males), and males from the Udzungwa Mountains (3.8– 4.5 mm) are thicker than all others (2.5–3.7 mm), except for the diameter of 4 mm for the type specimen of P. tersus published by Attems (1914).

The descriptions and illustrations of the very characteristic gonopods agree as good as completely between the various authors. Some apparent differences may be due to slight differences in the angle of view. For example, in the drawings by Mwabvu & VandenSpiegel (2009) the constriction at midlength of the lateroapical metaplical process (lap) is not so obvious, but this may be due to a slightly apical view.

The most deviating specimen is the topotype of Ptilostreptus tersus from Mpwapwa District, but with only a single specimen from Mpwapwa District at hand, the significance of these slight differences cannot be properly evaluated, and we hesitate to ascribe this specimen to a separate species.

Similarly, based on the confusing situation regarding L. regularis described above, there seems to be no justification for regarding L. regularis as a species distinct from L. tersus .

Figures

Fig. 1. Example of a millipede sample from the FoRCE experiment. Magombera Nature Park, Half- FoRCE Plot 38, 17 Feb. 2018. A.R. Marshall leg. 1. Lyodesmus dollfusi (Cook, 1896) (Oxydesmidae). 2. Lophostreptus tersus (Cook, 1896) (Spirostreptidae). 3. Tropostreptus sigmatospinus Enghoff, 2017 (Spirostreptidae). Photo: A.R. Marshall.

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. 14. Numbers of body rings in adults and epimorphic juveniles of Lophostreptus tersus (Cook, 1896) from the Udzungwa Mountains.

Fig. 15. Lophostreptus tersus (Cook, 1896), male from Udzungwa Mts National Park (NHMD 1184641). A–B. Head, collum and rings 2–6. A. Lateral view. B. Dorsal view. C–D. Midbody rings. C. Lateral view; inset: close-up of ozopore. D. Dorsal view; inset: close-up. E–G. Telson and last rings. E. Lateral view. F. Posterior view. G. Dorsal view. Abbreviations: av = anal valve; mz = metazonite; oz = ozopore; pr = preanal ring; pz = prozonite. Scale bars = 0.5 mm.

Fig. 16. Lophostreptus tersus (Cook, 1896), male from Udzungwa Mts National Park (NHMD 1184641), first pair of legs. A. Posterior view. B. Sublateral view. C. Ventral view. Abbreviations:aps = apicomesal prefemoral setae; cxs = coxosternal setae;pfl = prefemoral lobes. Scale bars: A = 0.2 mm; B–C = 0.1 mm.

Fig. 17. Lophostreptus tersus (Cook, 1896), male from Udzungwa Mts National Park (NHMD 1184641), gonopods. A–B. Coxae and left telopodite. A. Anterior view. B. Posterior view. C. Right telopodite, distal part. D–E. Two views of telopodite tip. Abbreviations: kn = knee; lap = lateroapical metaplical process; map = mesapical metaplical process; pal = palette-like appendage; slm = solenomere; tp1, tp2, tp3 = subapical telopodital processes. Scale bars: A–B = 0.2 mm; C = 0.1 mm; D–E = 0.01 mm.

Fig. 18. Lophostreptus tersus (Cook, 1896), specimens from the Udzungwa Mts, and L. magombera Enghoff sp. nov., number of body rings vs midbody vertical diameter. Notice that adult and juvenile epimorphic females have not been distinguished – the symbols in the lower range of the L. tersus female “cloud” certainly represent juvenile females.

Tables

Table 2. Characters of Lophostreptus tersus (Cook, 1896) according to previous authors and based on examined specimens.

Nominal species	Ptilostreptus tersus	Lophostreptus ptilostreptoides	Lophostreptus regularis	Lophostreptus malleolus		examined specimens of Lophostreptus tersus
Source(s)	Cook (1896); Attems (1914)	Carl (1909)	Attems (1909)	Kraus (1958)	specimens from Udzungwa Mountains	syntypes of L. regularis (NHRS-TOBI 000005480, NHRS-TOBI 000005478, NHRS-TOBI 000005476, NHRS-TOBI 000005481 NHRS-TOBI 0000077)	topotype of P. tersus (VMNH 112024)	specimens from Kimboza Forest Reserve (NHMD 1184657)	specimens from Arusha (NHMD 1184655)	specimen from Mindu Mountain (VMNH 112923)
Type locality	Mpapua	Bushland between Kagera and the Mabira camp in S Karagwe	Kilimanjaro: Kibonoto, steppe, cultural zone, mixed forest, 1000–1900 m • Massai steppe • Usambara: Tanga	Dar es Salaam, botanical garden	–	Kilimanjaro: Kibonoto, steppe, cultural zone, mixed forest, 1000–1900 m • Massai steppe • Usambara: Tanga	Mpapua (Mpwapwa)	–	–	–
No. of rings	47	45–48	46–49	49	48–52	46–48	51	49	48–50	47
	(48 segments)	(46–49 segments)	(47–50 segments)	(50 segments)
Male diameter	4 mm	3 mm	3 mm	3.0 mm	3.8–4.5 mm	3.0 mm	3.7 mm	3.4–3.7 mm	2.5–3.2 mm	3.4 mm
Pads on male legs	no	no	large pads on postfemur and tibia of pregonopodal legs	no	no	no	no	no	no	no