Henicops milledgei, Hollington & Edgecombe, 2004

Hollington, L. M. & Edgecombe, G. D., 2004, Two New Species of the Henicopid Centipede Henicops (Chilopoda: Lithobiomorpha) from Queensland and Victoria, With Revision of Species from Western Australia and a Synoptic Classification of Henicopidae, Records of the Australian Museum 56, pp. 1-28 : 16-22

Henicops milledgei n.sp.

Figs. 12–16

Etymology. For Graham Milledge, who collected this species throughout most of its range.

Type material. HOLOTYPE: MV NOH-2207, ♀ ( Fig. 12A), Strzelecki Ra , Tarra-Bulga NP, VIC, 0.5 km NNE Tarra Valley Picnic Area , 38°26'40"S 146°32'30"E, Nothofagus cunninghamii forest, G.A. Milledge, 14 Nov 1995 – 10 Jan 1996. P ARATYPES: NOH-2208, 1♀, 333, from type locality, 10 Jan–5 Mar 1996; NOH-2209, 1 ♀, 13, type locality, 5 Mar–7 Mar 1995; NOH-2211, 1 ♀, 13, Tarra-Bulga NP, 0.2 km W Tarra Valley Picnic Area, 38°27'S 146°32'E, 14 Sep–14 Nov 1995; NOH- 2212, 13, same locality, 14 Nov 1995 – 10 Jan 1996; NOH- 2213, 1 3, same locality, 14 Nov 1995 – 10 Jan 1996; NOH-2214, 1 ♀, 333, same locality, 10 Jan–5 Mar 1996; NOH-2215, 13, same locality, 7 May–16 Jul 1996; NOH-2220, 1♀, 233, same locality, 5 Mar–7 May 1996; NOH- 2217, 1♀, Tarra-Bulga NP, Bulga Picnic Area, 38°25'30"S 146°34'20"E, 14 Nov 1995 – 10 Jan 1996; NOH- 2218, 1 3, same locality, 10 Jan–5 Mar 1996; NOH-2219, 2♀♀, 13, same locality, 5 Mar–7 May 1996.

Other material. VICTORIA: MV: OTWAY RA, leg. G.A. Milledge: NOH-1045, 2♀♀, 13, Philips Track, 0.5 km N Triplet Falls , 38°40'S 143°29'E, 6 Sep–15 Nov 1994 ; NOH-1046, 1 ♀, 13, same locality, 15 Nov 1994 – 31 Jan 1995 ; NOH-1047, 3 ♀♀, same locality, 31 Jan–11 Apr 1995 ; NOH-1048, 13, same locality, 11 Apr–14 Jun 1995 ; NOH- 1049, 1♀, same locality, 14 Jun–29 Aug 1995 ; NOH-1050, 2 ♀♀, 233, Young Creek Rd, 0.4 km NW Triplet Falls , 38°40'S 143°29'E, 6 Sep–15 Nov 1994 ; NOH-1052, 7♀♀, 333, same locality, 15 Nov 1994 – 31 Jan 1995 ; NOH-1053, 1♀, same locality, 31 Jan–11 Apr 1995 ; NOH-1054, 3♀♀, same locality, 11 Apr–14 Jun 1995 ; NOH-1055, 3 ♀♀, 13, Young Creek Rd, 0.2 km NE Ciancio Ck Crossing, 38°40'S 143°29'E, 6 Sep–15 Nov 1994 ; NOH-1056, 1♀, same locality, 15 Nov 1994 – 31 Jan 1995 ; NOH- 1058, 2♀♀, 13, 31 Jan–11 Apr 1995; NOH-1059, 1 ♀ ( Figs. 14K,L, 15F, 16G–J), 2 33, same locality, 11 Apr–4 Jun 1995 ; NOH- 1060, 1 3, Aire Crossing Tk, 0.5 km N Aire R. Crossing, 38°40'S 143°29'E, 6 Sep–15 Nov 1994; NOH-1061, 233, Beauchamp Falls , 38°39'S 143°36'E, 6 Sep–15 Nov 1994 ; NOH-1062, 7 ♀♀, 13, same locality, 15 Nov 1994 – 31 Jan 1995 ; MV NOH-1063, 8♀♀, 13, same locality, 31 Jan–11 Apr 1995 .

CENTRAL HIGHLANDS, leg. G.A. Milledge: NOH-1778, 1 larval stadium LII, 2 larval stadia LIII, Cement Creek Reserve , 2.2 km ESE Mt Donna Buang, 37°43'S 144°42'15"E, 13 Dec 1994 ; NOH-1779, 2♀♀, 13 ( Figs. 14B,C,H–J, 15B,D,E,G,L,M, 16A,D–F,K,L), same locality, 29 Nov 1994 – 20 Jan 1995 ; NOH-1784, 3 ♀♀, same locality, 21 Jan–7 Apr 1995; MV NOH-1782, 4♀♀ ( Fig. 12B), 433, 1 larval stadium LII, Donna

Buang Rd, 1 km SW Mt Donna Buang , 37°43'S 144°40'E, 29 Nov 1994 – 20 Jan 1995 ; NOH-1783, 6♀♀, 13, same locality, 21 Jan–7 Apr 1996 ; NOH-1786, 2♀♀, 233, 1 larval stadium LII, Myrtle Gully Reserve , 3.4 km WSW Mt Donna Buang, 37°43'S 144°38'30"E, 29 Nov 1994 – 20 Jan 1995 ; NOH-1787, 5♀♀, 333, same locality, 21 Jan–7 Apr 1995 ; NOH-1788, 1♀, Road 26, 0.2 km WNW Donna Buang Rd junct., 37°43'S 144°39'30"E, 16 Feb 1995 ; NOH-1790, 1♀, Acheron Gap, 6 km NE Mt Donna Buang , 37°40'43"S 144°44'20"E, 20 Feb 1996 ; NOH-1792, 233 ( Fig. 12C), same locality, 26 Oct–28 Dec 1995 ; NOH-1793, 2 ♀♀, same locality, 28 Dec 1995 ; NOH-1794, 4♀♀, same locality, 21 Feb–23 Apr 1996 ; NOH-1795, 1♀, same locality, 23 Apr–25 Jun 1996 ; NOH-1796, 2♀♀, 233, 0.7 km N Acheron Gap , 7 km NE Mt Donna Buang, 37°40'17"S 144°44'20"E, 21 Feb–23 Apr 1996 ; NOH-1797, 4 33, same locality, 28 Dec 1995 – 21 Feb 1996 ; NOH-1800, 1 larval stadium LIII, The Big Culvert, 2.5 km ENE Mt Observation , 37°33'36"S 145°52'15"E, 28 Dec 1995 .

STRZELECKI RA, leg. G.A. Milledge: NOH-2222, 1 larval stadium LIII, Tarra-Bulga NP, Bulga Picnic Area , 38°25'30"S 146°34'20"E, 7 May– 16 Jul 1996; NOH-2225, 1♀ ( Fig. 13A–D), 2 33, Gunyah-Toora Rd, 2 km SSW Gunyah Gunyah, 38°32'30"S 146°19'E, 14 Nov 1995 –10 Jan

13, (C) leg 14, (D) leg 15. Scale 0.5 mm.

1996; NOH-2227, 1♀, Jeeralang West Rd , 0.1 km N Binns Hill Junction, 38°26'30"S 146°29'E, 10 Jan–5 Mar 1996 .

AUSTRALIAN MUSEUM: KS35864, 1♀, Otway Ra , 38°27'S 143°58'E, A. Frazer, Aug 1979 ; KS35869, 13, Otway Ra , Lavers Hill, 38°41'S 143°24'E, A.Frazer, 24 Feb 1979 ; KS83632, 1♀, Otway NP, Sandy Ridge , Spur Road, 38°46.12'S 143°32.9'E, 278 m, G. Cassis et al., 9 Nov 2002 ; KS83633, 2♀♀ ( Figs. 14A,D–G, 15A,C,H,K, 16B,C), Grampians NP, Wonderland Range , Silverband Rd, 1.2 km S Rosea Campground, 37°10'28"S 142°30'29"E, G.D. Edgecombe & Z. Johanson, 15 Feb 2000 , fern gully.

Distribution. Victoria: Otway Ranges, Strzelecki Ranges, Central Highlands, mostly from Nothofagus cunninghamii forest; Grampians, sclerophyll forest ( Fig. 1).

Diagnosis. Henicops with 36–43 (most commonly 40 or 41) antennal articles. Uneven change in length of articles in proximal part of antenna, with short paired articles interspersed between longer articles; T7 with rounded, concave margin; Tömösváry organ small and weak, centrally located on cephalic pleurite; groove in accessory denticle field pronounced on mandibular teeth; female gonopod with two moderately large, bullet-shaped spurs (shared with H. maculatus ). Mandibular aciculae differentiated into outer row with serrated margins and inner row with mostly simple margins; branching bristles of mandible with sparse spinelike branches on basal part of bristle (shared with H. dentatus Group). Uniquely: 3+3 moderate sized teeth on maxillipede coxosternum, with space between outer and middle teeth invariably more than twice the space between inner two teeth; three tarsomeres in leg 13, four in leg 14, five in leg 15, basitarsus undivided in legs 13–15; proximal coxal pores elliptical, distal pores round.

Description. Length up to 28 mm; width of head shield up to 3.3 mm. Colour (based on specimens in absolute ethanol): antenna dark orange, with dark pigment spots usually present, especially on dorsal side and proximally but also common on ventral side of antenna and more distally; head shield dark orange with chestnut mottled network; tergites dark orange with dark mottling covering most of tergite, concentrated in longitudinal median band and near margins, and dark pigment spots concentrated on margins; maxillipedes orange; sternites pale orange with purple mottling around margins, becoming more concentrated in the three posterior sternites; prefemur to tibia pale to moderate yellow with purple mottling; tarsi pale to moderate yellow; genital sternite and gonopods orange with faint purple mottling on genital sternite.

Head shield smooth. Frontal margin with strong median notch and short median furrow; posterior margin transverse; border very slightly wider posterolaterally than posteromedially. Antennal length 3.1–4.7 times width of head shield, usually extending back to tergite 5 ( Fig. 12A); 36– 43 articles, commonly 40 or 41; basal two articles much larger than succeeding two ( Fig. 14A); antenna with a markedly uneven change in the length of articles in proximal part, with short paired articles interspersed between longer articles ( Fig. 14B); articles mostly substantially longer than wide in distal half of antenna; antenna densely setose along most of length, with many short, curved sensilla interspersed with long trichoid sensilla ( Fig. 14C). Ocellus moderately large, overhanging lateral margin of head, usually translucent, whitish to dark purple, domed. Tömösváry organ small, fairly shallow, longitudinally elliptical, situated inward of anterolateral margin of cephalic pleurite ( Fig. 15D,E).

Clypeus with transverse band of six setae medially just in front of labrum ( Fig. 15B), cluster of 10 setae at clypeal apex ( Fig. 15A). Labral margin with rounded shoulder beside midpiece, with gradual break in curvature where fringe of branching bristles overhangs margin ( Fig. 15B); most bristles with short, spine-like projections along their length, some branching into long, slender spines distally ( Fig. 15C).

Maxillipede coxosternum trapezoidal to subsemicircular, dental margin broad, each half gently convex ( Fig. 16A,B); median notch at most narrow, shallow ( Fig. 16D); teeth small, triangular projections, invariably 3+3, with space between outer tooth and adjacent tooth more than twice space between inner two teeth ( Fig. 16C–E), inner tooth smaller than the others. Coxosternum bearing moderate number of small to moderate sized setae ( Fig. 16D), mostly confined to anterior half of coxosternum ( Fig. 16A,B). Tarsungulum with long, slender pretarsal section ( Fig. 16A,B). Setae on telopodite denser on inner part of femur, tibia and tarsal section of tarsungulum than on outer part, especially long on tarsungulum ( Fig. 16C).

Mandible as described for H. tropicanus except for the following: ventral aciculae in outer row of aciculae with both margins serrated along their distal fifth to third ( Fig. 14G,I); grooved ridge on accessory denticle field of all but ventralmost paired tooth ( Fig. 14D,F); ventral bristles in fringe of branching bristles lacking spine-like projections on basal fifth of bristles ( Fig. 14H); fringe of scale-like branching bristles extends to furry pad ( Fig. 14J); bristles of furry pad curved towards dorsalmost tooth ( Fig. 14F,J).

First maxilla ( Fig. 15H–J) as described for H. tropicanus except: coxal process with a few short simple setae, several laciniate and plumose setae with intermediates between these (some with a few terminal spines, others with numerous short, slender projections along their distal halves) ( Fig. 15J); distal article of telopodite with several simple setae scattered over its main field ( Fig. 15I). Second maxilla ( Fig. 15K–M) as described for H. tropicanus .

Tergites wrinkled, faintly so anteriorly but considerably so in posterior half of trunk. T1 trapeziform, about as wide as head shield ( Fig. 12A), about 80% width of widest tergite (T8), posterior angles rounded and posterior margin gently concave; lateral borders subparallel in TT3 and 5, posterior angles rounded, posterior margins gently concave; TT1, 3 and 5 bordered posteriorly; border of T7 incomplete posteriorly; posterior margin of T7 evenly concave, posterior angle blunt; posterior margins of TT8 moderately, fairly evenly concave, posterior angles obtuse, blunt; posterior margins of TT6, 11 and 13 all evenly concave, increasingly so more posteriorly; T9 variably with nearly transverse median sector; posterior angle of T6 rounded, TT9, 11 and 13 with triangular projections; posterior margin of T14 moderately concave, with obtuse posterior angles; tergite of intermediate segment with weakly concave posterior margin in female, moderately concave margin in male. Tergal surface and margins scattered with short setae, in some specimens relatively numerous, e.g., posterior margins of TT3 and 4 fringed by about 20 setae. Longitudinal median furrow confined to anterior part of sternites, not more than one-third length of sternite.

Legs: Distal spinose projections on tibiae strong, pointed on legs 1–14. Setae relatively uniformly distributed along length of legs, with similar setal density on inner and outer faces of all podomeres; legs 14 and 15 ( Fig. 13C,D) with similar setal density as 1–13 ( Fig. 13A,B). Three tarsomeres in legs 1–12 ( Fig. 13A), with divided basitarsus; tarsal articulations discontinuous on dorsal side of leg in anterior part of trunk; three pairs of consistently strong, pigmented, divergent setae on tarsi of legs 1–12, one just proximal to each articulation between tarsomeres, one near midlength of distal tarsomere. Basitarsus-distitarsus joint with a condyle in legs 13–15 only, basitarsus undivided in these three legs; three tarsomeres in leg 13 ( Figs. 13B, 15F); four tarsomeres in leg 14 ( Fig. 13C), distitarsus about 65% length of basitarsus, three tarsomeres comprise about 45–55, 15– 25 and 25–35% (proximal to distal) length of distitarsus; five tarsomeres in leg 15, the proximal distitarsal podomere about 50–55% length of distitarsus, second and third shorter than fourth ( Fig. 13D). Distitarsus 70–85% length of basitarsus in leg 15; basitarsus 10–12 times longer than broad. Pretarsus with anterior and posterior accessory claws on all legs, about 30% length of main claw, inserted on dorsolateral side of main claw, with gentle dorsoventral curvature ( Fig. 14K). Main claw gently curved, divided into many elongate, scales along most of its length, with short, polygonal scales ventrolaterally in region beneath proximal part of accessory claws, a few pores between these scales ( Fig. 14L), pore openings in the middle of small polygons; scales weakly defined proximally beneath bases of accessory claws and on proximoventral surface of main claw, well defined dorsoproximally. Posteroventral spine ('sensory spur' of Eason, 1964: fig. 486) present on legs 1– 14 ( Fig. 14L), absent on leg 15 ( Fig. 15G), about 15% length of main claw, directed distally, bearing slender subsidiary spine that parallels it on its dorsoproximal half ( Fig. 14L); accessory claws and posteroventral spine with surface ornament of linear grooves and ridges.

Coxal pores round, transversely ovate, or elliptical, typically with round distal pores and elliptical proximal pores ( Fig. 12B); overwhelmingly 6,5,5,5/6,5,5,5 in large females (width head shield>1.75 mm), 5,4,4,4/5,4,4,4 in large males; maximum in males 5,5,5,5/5,5,5,5. Coxal pore field set in shallow groove. Anal pores large in both sexes.

Female ( Fig. 12B): Sternite of segment 15 transverse or convex posteromedially, with about a dozen setae along posterolateral and posteromedial margins. Tergites of first genital segment and telson usually well sclerotized; sternite of first genital segment large, posterior margin concave, with median bulge, between condyles of gonopods, surface scattered with setae, longest and densest posterolaterally ( Fig. 16G). First article of gonopod bearing two moderately large, bullet-shaped spurs, inner spur slightly smaller ( Fig. 16G–J); spurs gently curved such that tip points up ( Fig.

16I); first article usually bearing more than 30 setae in large specimens (up to 38), second article with 12–20 moderately long setae; third article with three to eight setae. Claw large, undivided, numerous pores with sensilla coeloconica on dorsodistal surface of claw ( Fig. 16J).

Male ( Fig. 12C): Sternite of segment 15 with subtransverse ( Fig. 12C) to moderately convex ( Fig. 16F) posteromedial margin. Sternite of first genital segment small, divided, with transverse posterior margin, bearing about 20 moderate to long setae on each half with slight to moderate concentration near posterior margin. Gonopod of three articles and tapering, seta-like terminal process ( Fig. 16K,L), with many short spine-like projections along its inner margin; articles of gonopod each typically bear 10– 12 setae; many setae on gonopod and first genital sternite curved, at least distally.

Larval stadia: Stadium LII with 8 pairs of legs and two pairs of limb-buds; 14 antennal articles, all elongate; 3+3 teeth on dental margin of maxillipede coxosternum, outer tooth more distant than inner pair; tarsi unjointed. Stadium LIII with 10 pairs of legs and two pairs of limb-buds; 25 antennal articles; maxillipede dentition as in stadium LII; single tarsal articulation faintly defined on many legs.

Discussion. All specimens of Henicops milledgei share a large space between the outer and middle tooth of the maxillipede coxosternum (more than twice the distance between the inner and middle tooth), exceeded only by H. brevilabiatus . No notable geographic variation has been detected between populations of H. milledgei from different regions of Nothofagus forest in Victoria.

Segmentation of the larval antenna of Henicops milledgei and H. tropicanus presents a character that may be autapomorphic for Henicops or a group within the genus. Larval stadium LIII in both species has a uniquely high number of antennal articles (25), a count retained in larval stadium LIV in H. tropicanus at least. In comparison, Lamyctes emarginatus (= fulvicornis ) and L. coeculus have only 14 articles in stadium LIII, and Lithobiidae have 14– 17 articles in this stadium ( Andersson, 1979: table III). Of Lithobiomorpha for which larval ontogeny has been described, only Cermatobius longitarsis approaches Henicops in having 20–24 articles in stadium LIII ( Murakami, 1960; Andersson, 1979).