Solaenodolichopus pruvoti ( Brolemann, 1931 )

Mesibov, Robert, 2014, The millipede genus Solaenodolichopus Verhoeff, 1924 (Diplopoda, Polydesmida, Paradoxosomatidae). 1. New genus diagnosis and redescriptions of named species, European Journal of Taxonomy 83, pp. 1-36 : 11-17

Solaenodolichopus pruvoti ( Brolemann, 1931)

Figs 1F , 2A , 4A–B , 5 , 6A–B , 7 , 10A , 11D

Paraulacoporus Pruvoti Brolemann, 1931: 295 (first description); figs 29–32 (p. 297), 33–35 (p. 299). Solänodolichopus annulatus Verhoeff, 1941: 11 (first description); figs 1–4 (p. 23).

Solaenodolichopus pruvoti – Jeekel 2000: 40 (new combination), 42, 46. — Nguyen & Sierwald 2013: 1159.

Solaenodolichopus annulatus – Jeekel 2002: 60 (as synonym of Solaenodolichopus pruvoti ).

Solänodolichopus annulatus – Nguyen & Sierwald 2013: 1159 (as synonym of Solaenodolichopus pruvoti ).

Aulacoporus pruvoti – Attems 1937: 261 (new combination), 264 ( Brolemann’s 1931 description reworded); fig. 331 (p. 265; same as fig. 32 in Brolemann 1931). — Jeekel 1968: 18, 29; 1981: 47; 1985: 34.

Aulacoporus annulatus – Jeekel 1968: 20, 29; 1981: 49; 2000: 41 (synonymised with Solaenodolichopus pruvoti ), 42.

non? Solaenodolichopus pruvoti – Jeekel 2002: 60 (partial redescription from new material), 77; fig. 1 (p. 62).

non? Aulacoporus pruvoti – Jeekel 1982: 124 (redescription from new material); figs 2, 3 (p. 126).

Material examined

Syntypes of S. pruvoti

At least 1 male and 1 female, Dumbéa, New Caledonia [22°09’ S, 166°27’ E, ± 5 km], date unknown, A. Pruvot-Fol; specimens not located.

Syntypes of S. annulatus

At least 1 male, Brisbane, QLD [27°28’ S, 153°01’ E, ± 10 km], Dec. 1936, J. Mauritzon; specimens not located.

Other material

QUEENSLAND: 1 ♂, Mt Glorious [27°20’ S, 152°46’ E, ± 2 km], Oct. 1970, H. Williams, QM S5944; 1 ♂, Mt Nebo [27°23’ S, 152°47’ E, ± 2 km], 19 Nov. 1978, A. Rozefelds, pitfall, wet sclerophyll, QM S5945; 2 ♂♂, 1 ♀, Camira, 27°37’ S, 152°55’ E [± 2 km], 30 Jul.–Oct. 1990, R. Raven, open forest, pitfall, QM S19631 ; 1 ♂, Mt Glorious barracks, 27°19’54” S, 152°45’12” E [± 500 m], 660 m a.s.l., 7 Dec. 1991 – 6 Mar. 1992, G. Monteith, rainforest, pitfall+intercept, QM S74833 ; 1 ♂, Mt Mee, 27°02’53” S, 152°40’49” E [± 500 m], 300 m a.s.l., 3 Mar.–12 Apr. 1992, D. Cook, rainforest, intercept, QM S74834 ; 2 ♂♂, Stony Creek via Samford, 27°20’20” S, 152°47’52” E [± 500 m], 200 m a.s.l., 22 Oct. 1994 – 2 Feb. 1995, H. Janetzki and G. Monteith, rainforest, pitfall, QM S74835 ; 1 ♂, 1 ♀, Camerons Scrub, knoll top, 27°30’27” S, 152°43’40” E [± 500 m], 90 m a.s.l., 15 Sep.–11 Nov. 1998, G. Monteith, D. Cook and G. Thompson, vine scrub, pitfall, sample 7380, QM S74841 ; 3 ♂♂, 1 ♀, same details but 11 Nov. 1998 – 13 Jan. 1999, sample 7557, QM S74856 ; 1 ♂, same details but Camerons Scrub, top ridge, 27°30’29” S, 152°43’54” E [± 500 m], 160 m a.s.l., 21 Sep.–11 Nov. 1998, sample 7381, QM S74842 ; 1 ♂, 1 ♀, same details but 13 Jan.–16 May 1999, sample 7666, QM S74864 ; 1 ♂, 1 ♀, same details but Camerons Scrub, below road, 27°30’45” S, 152°43’39” E [± 500 m], 40 m a.s.l., 11 Nov. 1998 – 13 Jan. 1999, sample 7555, QM S74857 ; 1 ♂, The Knobby, 27°30’27” S, 152°35’18” E [± 500 m], 240 m a.s.l., 11 Nov. 1998 – 13 Jan. 1999, G. Monteith, D. Cook and G. Thompson, vine scrub, pitfall, sample 7567, QM S74858 ; 1 ♂, Split Yard Creek, 27°22’41” S, 152°37’56” E [± 500 m], 150 m a.s.l., 27 Dec. 1998 – 13 Jan. 1999, G. Monteith, D. Cook and G. Thompson, vine scrub, dung trap, sample 7568, QM S74859 ; 1 ♂, Boombana, Brisbane, 27°24’08” S, 152°47’22” E [± 100 m], 440 m a.s.l., 7 Nov. 2003, R. Raven, B. Baehr and O. Seeman, day hand collection, ex S.C. 51695, QM S74874 .

WESTERN AUSTRALIA: 1 ♂, 1 ♀, Curtin University, Environmental Biology Building, Bentley, [32°00’21” S, 115°53’40” E, ± 200 m], 25 Apr. 2002, T. Cocket, WAM T47890; 1 ♂, same details but Physics Building, 32°00’23” S, 115°53’45” E [± 50 m], 10 m a.s.l., 4 Nov. 2008, M.S. Harvey, in building, WAM T73493; 1 ♂, 1 ♀, Nursing Home, Hamersley Road, Subiaco [31°57’28” S, 115°49’25” E, ± 100 m], 30 m a.s.l., 22 Jan. 2003, K. Debnam, inside nursing home, WAM T47891; 1 ♀, 7 Kingston Street, Nedlands, 31°57’56” S, 115°48’46” E [± 100 m], 10 m a.s.l., 19 Apr. 2003, J.M. Waldock, found dead on front verge near driveway, WAM T52005; 1 ♂, same details but 21 Apr. 2003, J.M. Waldock, walking on footpath in front garden, WAM T52006; 1 ♂, same details but 19 May 2003, S. Slack-Smith, in garden, WAM T52345; 1 ♂, same details but 28 Sep. 2007, WAM T95077; 2 ♀♀, same details but 6–9 Nov. 2008, WAM T94065; 2 ♂♂, 1 juvenile, same details but 6 Nov. 2008, on path in garden after rain, WAM T94986; 2 ♀♀, 3 juveniles, same details but 8 Dec. 2008, on path in garden, WAM T94973; 1 ♂, 22 Windelya Road, Murdoch [32°04’14” S, 115°49’31” E, ± 50 m], 30 m a.s.l., 3 Apr. 2005, O. Mueller, WAM T63211; 1 ♂, same details but 24 Apr. 2005, on verandah at night, WAM T65825; 1 ♀, Edward Street, Nedlands, 31°58’ S, 115°48’ E [± 2km], 20 m a.s.l., 1 Mar. 2006, A. Baynes, WAM T67503; 2 ♂♂, Garfield Way, Greenwood, 31°49’ S, 115°48’ E [± 2 km], 40 m a.s.l., 25 May 2006, T. Struthers, garden litter, WAM T78133; 1 ♂, 3 ♀♀, same details, WAM T112662; 1 ♂, 56 Edward Street, Nedlands [31°58’54” S, 115°48’24” E ± 50 m], 20 m a.s.l., 3 May 2007, A. Baynes, WAM T82708; 1 ♂, University of Western Australia, Crawley, bus stop on Fairway near corner of Cooper Street, 31°58’41” S, 115°48’56” E [± 100 m], 11 Jun. 2008, J. Wojcieszek, WAM T89288; 1 ♂, 7 Dunkeld Street, Floreat, 31°55’51” S, 115°47’24” E [± 100 m], 13 Dec. 2012, J. Foss, WAM T129488.

Description

(Based mainly on Camerons Scrub, QLD specimens.) As for the genus. Maximum male/female midbody width ca. 3.8/4.4 mm. Colour in alcohol: “Colour brown-black, ringed tawny-pink on the anterior (nested) end of the prozonites and on the median portion of the metazonites. Anal valves black. Legs pale yellow” ( Brolemann 1931: 295, my translation); “Bright gray-yellowish with brown posterior margin rings on diplosegments, head mainly dark brown” ( Verhoeff 1941, p. 11, my translation); base colour in QLD and WA specimens varying from pale yellow to light brown, with narrow, dark brown ring at rear of haplosegments, diplosegments and apodous ring; darker specimens with very narrow, light brown rings bordering waist; narrow marginal ring of dark brown on collum, brown patches on vertex and clypeus of head and on anal valves, antennomeres brown (darker distally), legs yellowish ( Fig. 4A–B ).

Male with transverse furrows very shallow, narrow, sometimes indistinct and easily overlooked (“Pas de sillon tranverse sur les métazonites”, Brolemann 1931: 296; “Querfurchen sehr fein”, Verhoeff 1941: 10). No longitudinal furrows laterally on diplosegments ( Fig. 4B ). Sternal lamella ( Fig. 2A ) with sides more or less straight, corners broadly rounded, distal margin convex. Scopulae on legs 1 to 29, i.e. not present on last podous ring. Leg bases on rings 6 and 8 separated more widely than on other rings, sternites a little depressed. Anterior margin of aperture with rounded-triangular, medial extension and shorter, gently convex extension on either side (as shown in Fig. 3B for S. rubriventris ).

Gonocoxa ca. 1/2 telopodite length, slightly flattened anteroposteriorly and slightly concave anteriorly, with sparse, long setae anterodistally. Cannula short, narrow, uniformly tapering towards prefemur. Gonopod telopodite ( Figs 5A–B , 6A–B , 7 , 10A ) reaching leg 4 bases on uncoiled specimens. Prefemur small, rounded, densely setose posteromedially, demarcated from femorite laterally by small, narrow notch. Remainder of telopodite clearly divided into femorite and solenomere at ca. 1/2 telopodite length. Femorite slender, straight in posterior view, slightly convex anteriorly in lateral view, the posterior surface very slightly concave; tipped with knob-like, rounded shoulder just lateral to lateral femorite process, a short, wide, longitudinal groove running basally from shoulder. Lateral femorite process a short, rounded tab directed anterodistally, close to solenomere base, extending distally a little past femorite shoulder. Medial femorite process teardrop-shaped, the rounded base with thin, translucent cuticle, the point directed anterodistally, terminating close to solenomere at 1/4-1/3 solenomere length. Solenomere at base ca. 2/3 femorite width, expanded and anteroposteriorly flattened distally; a narrow, ridge-like flange (arrow in Fig. 7D ) along whole of anterior surface of solenomere, diverting medially in short loop at ca. 1/2 solenomere length, curving sharply laterally at base to form small, rounded, anterior extension. Solenomere divided at ca. 2/3 length by large, U-shaped indentation into medially directed apex and distomedially directed, rounded process terminating just basal to solenomere apex; apex divided by narrow indentation into anteroposteriorly flattened, medially directed, truncate, subapical tab carrying opening of prostatic groove, and apical, basodistally flattened tab extending medially just past subapical tab. Prostatic groove ( Fig. 7D ) running on anterior surface of telopodite, following curve of solenomere medial to flange, terminating at midpoint on medial margin of subapical tab.

Female with leg 2 coxa extended posteroventrally as apically rounded process ( Fig. 5 C–D).

Distribution

Known from open forest, rainforest and vine scrub to at least 660 m a.s.l. in the D’Aguilar Range and adjoining areas west of Brisbane in southeast Queensland, from Mt Mee south to Ipswich, a northsouth extent of ca. 65 km (map Fig. 12 ). This is almost certainly the native range of the species, as very similar, undescribed species are found elsewhere in southeast Queensland. Introduced and established in the Perth metropolitan area in Western Australia; so far reported from Kingsley south to Kardinya, a north-south extent of ca. 30 km. Introduced (and established?) in New Caledonia. Overlaps in range with S. vittatus .

Taxonomic notes

The types of P. pruvoti and S. annulatus have not yet been located, and I prefer not to designate lectotypes for these species based only on the published descriptions and illustrations of Brolemann (1931) and Verhoeff (1941). The Brisbane and Perth specimens listed above agree closely with those descriptions and illustrations ( Figs 6 A–B, 7A–B).

I suspect that the South Australian specimens described by Jeekel (1982, 2002) are an unnamed Solaenodolichopus species, not S. pruvoti . They differ from S. pruvoti in lacking pleural keels and in having a distinct transverse furrow on the metazonites ( Jeekel 1982: 125) and a medial coxal process on male legs 6 and 7 ( Jeekel 1982: 127). The specimens described in 2002 also differ from S. pruvoti in lacking dark transverse annulation ( Jeekel 2002: 61). Both Brolemann (1931: 295, for S. pruvoti ) and Verhoeff (1941: 11, for S. annulatus ) note strongly contrasting annular rings of colour, and all non-type adults listed above are clearly ringed ( Fig. 4A–B ), apart from a decoloured male in QM S74859 . Jeekel explained the difference in shape of the solenomere between S. pruvoti and the South Australian males ( Fig. 6 ) by saying “It should be emphasised that the distal portion of the gonopod (i.e. the solenomerite) is quite flexible, and may be curving more or less widely in a caudal direction, and may be crooked at the base” ( Jeekel 1982: 127). It is true that the S. pruvoti solenomere can be made to curve posteriorly by manipulation. However, in none of the preserved specimens I have so far examined of any Solaenodolichopus species is the solenomere ‘naturally’ curved as sharply as illustrated by Jeekel ( Fig. 6C ).

General notes

The collector of S. annulatus, Johan Mauritzon (1902-1950), was a lecturer in botany at Lund University when he travelled to Australia in 1936 in company with University of Uppsala zoologist Folke Linder (as reported in several contemporary Australian newspapers, e.g., The Adelaide Chronicle, 10 Sep. 1936: 52, http://trove.nla.gov.au/ndp/del/article/92463218, accessed 10 Oct. 2013). Mauritzon left Lund University soon after returning from his Australian trip (H. Wittzell, in litt. 10 Oct. 2013) and his Australian millipedes were later sent to Verhoeff from Lund by Torsten Gislén ( Verhoeff 1941: 3). The types of S. annulatus and four other species described by Verhoeff (1941) from this material have not yet been located. The types may not have been returned to Lund, as none of the Mauritzon millipede specimens collected in Australia are currently in the Lund University Zoological Museum (J. Ekstrom, in litt. 13 Jan. 2014).

The reported type localities of two of Verhoeff’s five 1941 species are incorrect, and it is unclear at which stage in the information chain from Mauritzon to Verhoeff the error appeared. Hoplatessara (Walestessara) forceps Verhoeff, 1941 (now Cladethosoma forceps ) and Hoplatria clavigera Verhoeff, 1941 were both described from Mauritzon collections in Gippsland, a district in eastern Victoria. Both have since been collected in NSW west of Sydney, and neither has since been found in intensively collected parts of Gippsland. However, the ‘Brisbane’ type locality for S. annulatus is plausible, because Verhoeff (1941: 12) gives “XII.36” as the collecting date and there are “near Brisbane” locations for Mauritzon plant collections dated 22 Dec. 1936 in the NHRS Herbarium (http://www.nrm.se/english/ researchandcollections/collections/databases/kryptos.8598_en.html, accessed 12 Oct. 2013) and in the Uppsala Museum of Evolution (http://130.238.83.220/botanik/home.php, accessed 12 Oct. 2013).

Brolemann’s 1931 paper is devoted to species collected by ‘Mme. Pruvot’, i.e., the malacologist Alice Pruvot-Fol (1873-1972), but Brolemann explicitly named Paraulacoporus pruvoti for Pruvot-Fol’s late husband, the zoologist Georges Pruvot (1852-1924) ( Brolemann 1931: 298).

Figures

Fig. 1. — A–B. Solaenodolichopus walesius Verhoeff, 1928. A. Lectotype ♂, AM KS.76508, right anterior view. B. ♂ ex AM KS.16152, midbody spiracles, ventral view, anterior to left. — C, E. S. vittatus (Verhoeff, 1924). C. Paralectotype ♀, NHRS KAS1000000007, midbody spiracles, ventral view, anterior to left (image right-left reversed for ease of comparison with 1B). E. ♂ ex QM S74828, anterior leg tip with scopula, tarsus at left. — D. S. teres (Verhoeff, 1924), lectotype ♂ slide mount, NHRS KAS1000000005, legpair 1. — F. S. pruvoti (Brolemann, 1931), ♂ ex QM S74856, epiproct, posterior and slightly ventral view. Scale bars: A = 2.5 mm, B–C = 0.5 mm, D = 1 mm, E = 0.25 mm, F = 0.1 mm.

Fig. 2. Posterior views of sternal lamella in ♂♂. — A. Solaenodolichopus pruvoti (Brolemann, 1931), QM S74856. — B. S. rubriventris Verhoeff, 1928, QM S74690. — C. S. sulcatus (Verhoeff, 1928), lectotype, AM KS.76503. — D. S. teres (Verhoeff, 1924), lectotype in alcohol, ZSM A20052188. — E. S. vittatus (Verhoeff, 1924), QM S74829. — F. S. walesius Verhoeff, 1928, lectotype, AM KS.76508. Scale bars: 0.5 mm.

Fig. 4. — A–B. Solaenodolichopus pruvoti (Brolemann, 1931), QM S74856. Dorsal (A) and lateral (B) views of midbody rings of ♂. — C–I. S. vittatus (Verhoeff, 1924). Dorsal (C-F) and lateral (G-I) views of midbody rings of ♂; QM S74767 (C, H), QM S74855 (D), QM S74836 (E, G), QM S74876 (F), QM S5946 (I). Scale bars: 2 mm.

Fig. 5. Solaenodolichopus pruvoti (Brolemann, 1931). A. Anterior lateral view showing reach of gonopod telopodites, QM S74856. B. Posterior view of gonopods, WAM T52345. C–D. Lateral (C) and posterior (D) views of leg 2 coxal process in ♀, QM S74857. Scale bars: A–B = 2 mm, C–D = 1 mm.

Fig. 6. A–B. Solaenodolichopus pruvoti (Brolemann, 1931); left gonopod solenomere, posterior views. A. Syntype, from fig. 32 in Brolemann (1931). B. QM S74841. — C. Solaenodolichopus?pruvoti; left gonopod solenomere, posterolateral view, from fig. 3 in Jeekel (1982). Scale bar: 1 mm.

Fig. 7. Solaenodolichopus pruvoti (Brolemann, 1931), left gonopod solenomere. — A–B. Medial views, S. annulatus (= S. pruvoti) syntype from fig. 2 in Verhoeff (1941) (A) and QM S74856 (B). — C–D. Lateral (C) and anterior (D) views, QM S74857. In D, arrow points to flange on anterior surface and dotted line marks course of prostatic groove. Scale bar: 1 mm.

Fig. 10. A. — Solaenodolichopus pruvoti (Brolemann, 1931), posterior view of gonopods in situ, QM S74858. — B–G. S. vittatus (Verhoeff, 1924). B–D, posterior view of gonopods in situ; QM S74826 (B), QM S95213 (C), QM S74767 (D). E–G, views of slide-mounts; left (E) and right gonopod solenomere (F) of S. vittatus lectotype, NHRS KAS1000000004, and left (top left) and right (bottom right) gonopod solenomere (G) of S. vittatus dorsalis holotype, NHRS KAS1000000003. Scale bars: A–D = 2 mm, E–G = 1 mm.

Fig. 11. — A–C, E–J. Solaenodolichopus walesius Verhoeff, 1928. A–B. Posterior views of gonopods in situ of lectotype, AM KS.76508 (A) and Parwalesoma castaneum Verhoeff, 1937 (= S. walesius) lectotype, ZMB 12652 (B). C. Posterior view of left gonopod solenomere tip, AM KS.16152. E–G, I. Lateral (E), anterior (F), medial (G) and posteromedial (I) views of left gonopod solenomere, AM KS.93755. H. Medial view of left gonopod solenomere of P. castaneum (= S. walesius), from fig. 6 in Verhoeff (1937). J. Posterior and slightly oblique view of left gonopod solenomere of S. walesius, from fig. 26 in Verhoeff (1928). — D. S. pruvoti (Brolemann, 1931), posterior view of left gonopod solenomere tip, QM S74856 (composite image at two focus levels). Dotted line in F marks course of prostatic groove. Scale bars: 1 mm.

Fig. 3. — A–B. Solaenodolichopus rubriventris Verhoeff, 1928, ♂, QM S74690. A. Right ventrolateral view of ring 7 showing swelling (arrow) just anterolateral to aperture. B. Ventral view of aperture showing gently convex extensions (arrows) on either side of medial extension on anterior margin of aperture. — C. S. walesius Verhoeff, 1928, ♀, AM KS.16152, posteroventral view of epigyne showing raised end (arrow). — D. S. teres (Verhoeff, 1924), paralectotype ♀, NHRS KAS1000000007, left ventrolateral view of anterior rings showing distolateral process on leg 2 coxa (arrow) just medial to raised end of epigyne. Scale bars: 1 mm.

Fig. 12. Localities for named Solaenodolichopus species as of February 2014. — A. Australia showing location of map B (rectangle). — B. S. teres (Verhoeff, 1924) (star), S. sulcatus (Verhoeff, 1928) (crosses) and S. walesius Verhoeff, 1928 (open triangles), other species (filled squares); rectangle shows location of map C. — C. S. pruvoti (Brolemann, 1931) (open squares), S. rubriventris Verhoeff, 1928 (crosses) and S. vittatus (Verhoeff, 1924) (open triangles). Maps A–C are geographic projections; approximate north-south map extents are 900 km for B and 220 km for C.