Heterolepisma buntonorum, Smith, 2016

Smith, Graeme B., 2016, On some Silverfish Taxa from Tasmania (Zygentoma: Lepismatidae and Nicoletiidae), Records of the Australian Museum 68 (2), pp. 45-80 : 58-65

Heterolepisma buntonorum n.sp.

Figs 79–120

Type material. Holotype ♀ ( HW 1.38 ) (K261010, K261011 on two slides) TAS: Bicheno lookout, 41.87731°S 148.30553°E, 30 m asl, 19.xii.2011, G. Smith and S. Bunton, under clean bark of live tree, three layers down . Paratypes (8♂♂, 4♀♀): ♀ ( HW 1.26 ) (K 377619 in alcohol) same data as holotype ; ♀ ( HW 1.18 ) (K 377620 in alcohol) same data as holotype ; ♂ ( HW 1.25 ) (K261012 K261013 on two slides) TAS: Bicheno lookout, 41.87757°S 148.30612°E, 64 m asl, 19.xii.2011, G. Smith and S. Bunton, leaf litter under rock overhang ; ♂ ( HW 1.00) (K 377621 in alcohol) same data as previous ; ♀ ( HW 1.20 ) (K 377622 in alcohol) same data as previous ;? ♂ ( HW 1.10 ) (K 377623 in alcohol, end of abdomen damaged) same data as previous ; juvenile ♀ ( HW 0.85) (K 377624 in alcohol) same data as previous ; juvenile ♂ ( HW 0.79) (K 377625 in alcohol) same data as previous ; juvenile ♂ ( HW 0.80) (K 377626 in alcohol) same data as previous ; ♂ ( HW 1.15 ) (gbs001772) used for scanning electron microscopy, same data as previous ; ♂ ( HW 1.13 ) (K 377627 in alcohol) TAS: Bicheno lookout, 41.87766°S 148.30613°E, 68 m asl, 19.xii.2011, G. Smith and S. Bunton, under bark of dead tree, possibly in abandoned termite workings ; ♂ ( HW 1.04 ) (K 377628 in alcohol) same data as previous .

Other material examined (2♂♂ 52♀♀): ♂ ( HW 1.05 ) (K261108 K261109 on two slides) TAS: Spring Beach , 42.58012°S 147.89847°E, 53 m asl, 19.xii.2011, G. Smith and S. Bunton, under bark of fallen Eucalypt in termite galleries ; ♂ ( HW 1.00) (K 377629 in alcohol), same data as previous ; juvenile ♀ ( HW 0.93) (K 377630 in alcohol) same data as previous ; juvenile ♀ ( HW 0.95) (K 377631 in alcohol) same data as previous ; ♀ ( HW 1.48 ) (K261014, K261015 on two slides) TAS: Friendly Beaches , 41.99301°S 148.27914°E, 63 m asl, 20.xii.2011, G. Smith and S. Bunton, dry woodland, bark spray to trunk of live Eucalypt ; ♀ ( HW 1.28 ) (K261016, K261017 on two slides) same data as previous ; ♀ ( HW 1.13 ) (K 377632 in alcohol) same data as previous ; ♀ ( HW 1.05 ) (K 377633 in alcohol) same data as previous ; ♀ ( HW 1.43 ) (K 377634 in alcohol) TAS: Friendly Beaches , 41.99296°S 148.27901°E, 63 m asl, 20.xii.2011, G. Smith and S. Bunton, dry woodland, pyrethrum spray into burned hollow in Eucalypt ; ♀ ( HW 1.05 ) (K 377635 in alcohol) same data as previous ; juvenile ♀ ( HW 0.85) (K 377636 in alcohol) same data as previous ; juvenile ♀ ( HW 0.58) (K 377637 in alcohol) same data as previous ; ♀ ( HW 1.28 ) ( TMAG F 14810 in alcohol) same data as previous ; ♀ ( HW 1.30 ) ( TMAG F 14810 in alcohol) same data as previous ; ♀ ( HW 1.20 ) (K 377638 in alcohol) TAS: Friendly Beaches , 41.99307°S 148.27903°E, 62 m asl, 20.xii.2011, G. Smith and S. Bunton, dry woodland, pyrethrum spray fallen Eucalyptus ; ♀ ( HW 1.00) (K 377639 in alcohol) same data as previous ; ♀ ( HW 0.93) (K 377640 in alcohol) same data as previous ; juvenile ♀ ( HW 0.83) (K 377641 in alcohol) same data as previous ; juvenile ♀ ( HW 0.88) (K 377642 in alcohol) same data as previous ; juvenile ♀ ( HW 0.78) (K 377643 in alcohol) same data as previous ; juvenile ♀ ( HW 0.85) (K 377644 in alcohol) same data as previous ; juvenile ♀ ( HW 0.85) (K 377645 in alcohol) same data as previous ; juvenile ♀ ( HW 0.80) (K 377646 in alcohol) same data as previous ; juvenile ♀ ( HW 0.78) (K 377647 in alcohol) same data as previous ; ♀ ( HW 1.30 ) (K261112 K261113 on two slides) TAS: Douglass-Apsley NP, Apsley Gorge near car park, 41.86514°S 148.18968°S 102 m asl, 20.xii.2011, G. Smith and S. Bunton, pyrethrum spray to bark of Eucalypt (with ants of the genus Ochetellus Shattuck, 1992 ); juvenile ♀ ( HW 0.83) (K 377648 in alcohol) same data as previous ; ♀ ( HW 1.49 ) (K 377649 in alcohol) same data as previous ; ♀ ( HW 1.30 ) (K 377650 in alcohol) same data as previous ; ♀ ( HW 1.33 ) (K 377651 in alcohol) same data as previous ; ♀ ( HW 1.23 ) (K 377652 in alcohol) same data as previous ; juvenile ♀ ( HW 0.95) (K 377653 in alcohol) same data as previous ; juvenile ♀ ( HW 0.98) (K 377654 in alcohol) same data as previous ; juvenile ♀ ( HW 0.80) (K 377655 in alcohol) TAS: Douglass-Apsley NP, Apsley Gorge nr. car park, 41.86509°S 148.18993°E, 111 m asl, 20.xii.2011, G. Smith and S. Bunton, pyrethrum spray to end of rotting log ; ♀ ( HW 1.30 ) (K261114 K261115 on two slides) TAS: Wyemans River-Bluemans Creek State Reserve , 42.01506°S 147.95835°E, 322 m asl, 21.xii.2011, G. Smith and S. Bunton, under bark of Eucalypt ; ♀ ( HW 1.20 ) (K 377656 in alcohol) same data as previous ; ♀ ( HW 1.13 ) (K 377657 in alcohol) same data as previous ; juvenile ♀ ( HW 0.95) (K 377658 in alcohol) same data as previous ; juvenile ♀ ( HW 0.90) (K 377659 in alcohol) same data as previous ; ♀ ( HW 1.30 ) (K261116 K261117 on two slides) TAS: High point on road between Bicheno and Campbell Town, 42.02539°S 147.76409°E, 629 m asl, 21.xii.2011, G. Smith and S. Bunton, under bark of Eucalytpus obliqua L’Her .; ♀ ( HW 1.33 ) (K 377660 in alcohol) same data as previous ; ♀ ( HW 1.50 ) (K 377661 in alcohol) same data as previous ; ♀ ( HW 1.23 ) (K 377662 in alcohol) same data as previous ; ♀ ( HW 1.20 ) (K 377663 in alcohol) same data as previous ; ♀ ( HW 1.13 ) (K 377664 in alcohol) same data as previous ; ♀ ( HW 1.00) (K 377665 in alcohol) same data as previous ; juvenile ♀ ( HW 0.90) (K 377666 in alcohol) same data as previous ; juvenile ♀ ( HW 0.95) (K 377667 in alcohol) TAS:above St Peters Pass rest area, 42.24176°S 147.40593°E, 373 m asl, 21.xii.2011, G. Smith and S. Bunton, under bark; juvenile ♀ ( HW 0.68) (K 377668 in alcohol) same data as previous ; ♀ ( HW 1.33 ) (K261110 K261111 on two slides) TAS: Mt Stuart lookout, 42.87437°S 147.29574°E, 247 m asl, 22.xii.2011, G. Smith and S. Bunton, under bark of dead tree ; ♀ ( HW 1.25 ) (K 377669 in alcohol) same data as previous ; ♀ ( HW 1.10 ) (K 377670 in alcohol) same data as previous ; ♀ ( HW 1.28 ) (K 377671 in alcohol) same data as previous , ♀ ( HW 1.05 ) (K 377672 in alcohol) same data as previous ; ♀ ( HW 1.10 ) (K 377673 in alcohol) same data as previous ; juvenile ♀ ( HW 1.00) (K 377674 in alcohol) TAS: Steppes (central highlands), 42.11°S 146.89°E, 844 m asl, 1.vii.2012, S. Bunton .

Diagnosis. No other adequately described species of Heterolepisma is reported to have 3+3 combs on urotergite I, urosternites II–VIII with 1+1 single macrochaetae and two pairs of styli in the ♀ but only one pair in the ♂.

Description. Appearance. Medium to large silverfish, scale covering in life uniform or slightly mottled grey with light brownish antennae, nota with lighter lateral margins, terminal filaments distinctly banded ( Fig. 79 ).

Body length. H+B up to 10.25 mm (♀) 8.65 mm (♂); maximum HW 1.50 mm; thorax: length up to 3.45 mm (or 0.28–0.35 H+B); width up to 2.20 mm with no great difference between the pro, meso- and metanota; maximum preserved length of antenna 6.3 mm (or 0.69 H+B); terminal filaments damaged in most specimens, maximum preserved length of apparently intact cercus 6.13 mm (or 0.76 H+B); maximum preserved length of median dorsal appendage 9.30 mm (or 0.95 H+B). Body neither elongate nor broad ( Fig. 80 ) with thorax slightly wider than abdominal segment I, the following abdominal segments about the same width until the fourth or fifth after which it tapers posteriorly.

Pigmentation. Pigment brown in alcohol preserved specimens but fades somewhat in slide mounted material. Pigment laterally on head especially behind the antennae and surrounding the eyes with bands across the frons between the antennae, the distal half of clypeus and labrum; pedicel and scape quite darkly pigmented, rest of flagellum uniformly lightly pigmented; all articles of maxillary palp strongly pigmented with denser pigmentation of articles two and three but less on the ultimate article especially apically; all articles of labial palp with strong pigmentation except for apex and distal half of face bearing papillae; pronotum with pigmentation along anterior margin and anterolateral corners as well as along lateral margins, meso- and metanota pigmented laterally; legs with pigmentation along outer edge of precoxa, along the length of the outer margin and distally on the inner margin of the coxa, trochanter with patch on margin distally, femur completely pigmented and darkest along inner margin and at apex of outer margin, tibia and first tarsal article evenly darkly pigmented with pigment becoming lighter on more distal tarsal articles, urotergite X dark around margins, coxites IX and styli IX pigmented, divisions of terminal filaments dark except for a distinctly unpigmented annulus at the distal end of each division around the rosette of large macrochaetae. Some individuals show greater or lesser levels of pigmentation, with less pigmentation in juvenile specimens.

Macrochaetae. Bifid apically, with longitudinal ribs and grooves when examined with SEM ( Fig. 92 ) or smooth, hyaline or brown ( Fig. 81 ) when examined with light microscope.

Scales. Unevenly rounded or ovoid, with numerous parallel ribs, that do not extend beyond the margin; in alcohol dorsal scales and the more lateral scales of the urosternites with dark brown ribs (much darker in their apical half) ( Fig. 82 ); ventrally mostly hyaline. Lanceolate scales were not observed.

Head. Wider than long and with chaetotaxy typical of the genus ( Fig. 83 ) i.e. marginal rows about two macrochaetae wide along the sides of the vertex, and a complete anterior row (i.e. no gap in the middle), the lateral rows extending back along the margin to the eyes and extending as a single short row of about six macrochaetae above the eyes and also below the eyes, as well as a small group extending sub-perpendicular to the margin at the level of each antenna; clypeus with numerous setae, some long and thin, and 1+1 combs of 4–6 macrochaetae proximally at the lateral ends of the suture with the frons; labrum with many long thin setae. Scales on top of head only. Eyes dark. —Antennae ( Figs 84–86 ) long, the more apical intervals with rare or inconspicuous rod-like basiconic sensillae near the apex of each annulus ( Fig. 93 ) as well as a small trichobothrium subapically on each interval. —Mandibles ( Figs 87, 88 ) typical for genus with well-developed molar and incisor areas; a group of about eleven strong and six finer apically bifurcated setae distally adjacent to the pectinate molar area and a bush of 40+ setae and macrochaetae externally. —Maxilla ( Figs 89, 90 ) with three large macrochaetae externally proximal to the palp, the lacinia with three strong teeth, one shorter than the rest, seven lamellate processes and a row of seven apically bifurcate setae, apical article of maxillary palp 1.5–3.6 times longer than wide and 0.8–1.7 times longer than penultimate article, the ultimate article in both sexes with three “branched” papillae. —Labium ( Fig. 91 ) short and broad with rows of strong bifurcated setae on the prementum, submentum with numerous long thin setae as well as 2+2 stronger setae laterally near the suture with the prementum, glossae and paraglossae quite broad with short curved setulae; labial palp short, apical article eccentric suboval, about as long as wide (L/W 1.0–1.2) with 2+3 papillae of compact type ( Fig. 94 ) and at least one curved club-like thin-walled basiconic sensilla.

Thorax. Pronotum ( Figs 96, 97 ) with wide setal collar of shorter and longer, apically bifurcated setae and macrochaetae as well as cilia, not arranged in distinct rows but about two or three macrochaetae wide; lateral margins also with numerous shorter and longer setae and some cilia, each side with several larger submarginal macrochaetae; trichobothrial areas open and in contact with the lateral margins, the anterior one slightly posterior to the mid-point along the margin usually with one large macrochaeta located mediad to the trichobothrium and several setulae (in one specimen K261012 the large macrochaetae are absent on both sides, on another K261114 it is missing on the right side but present on the left, and vice versa on specimen K261112); posterior trichobothrial area near posterior lateral corner with two submarginal macrochaetae as well as several setulae and a few cilia, the trichobothrium located in the most mediad position; posterior margin slightly concave with 1+1 combs each of one macrochaeta with a smaller seta between it and the margin and 1–3 cilia and/ or setulae. —Mesonotum ( Figs 98, 99 ) lacking anterior notal collar, lateral chaetotaxy similar to pronotum with the submarginal macrochaetae single in most cases, although in some specimens (e.g., K261012) one comb or two combs (K261116) are composed of two macrochaetae, the anterior trichobothrial areas located more posteriorly at about ⅔ of the distance along the margin, with the trichobothrium located between the macrochaeta and the margin and slightly anterior to the macrochaeta. —Metanotum ( Figs 100, 101 ) similar to mesonotum except posterior margin slightly more concave and the submarginal combs in the middle of the lateral margin are more likely to be composed of two macrochaetae (although on ♀ K261014 the comb of two macrochaetae is only present on the left side and another ♀ K261016 only on the right side anterior to the midpoint); the two most anterior submarginal macrochaetae may also lie quite close together almost appearing as another comb.

Presternum narrow, with transverse row of strong setae. —All thoracic sterna with hyaline scales. Prothoracic sternum large, only slightly longer than wide at its base and reaching almost to the end of the coxa, rounded apically ( Figs 102, 103 ), full length of lateral margins with numerous small marginal setae and cilia, 1+1 larger apically bifurcate macrochaetae distally near the margin as well as another three or four smaller submarginal macrochaetae on each side in the distal third, distinct combs absent. —Mesosternum ( Fig. 104 ) slightly longer than broad with marginal setae and cilia in distal half as well as 2+2 smaller subdistal macrochaetae and 1+1 larger macrochaetae slightly anterior to these. —Metasternum ( Fig. 105 ) more rounded, distinctly wider than long with marginal setae and cilia along the distal half of the lateral margins, 1+1 longer subapical simple macrochaetae between which the margin is largely glabrous and two or three subdistal submarginal apically bifurcate macrochaetae.

Legs fairly long ( Figs 102, 106 ), tibia L/W ratio of legs PI 2.5–3.4, PII 2.6–3.8, PIII 2.4–5.0; tarsi L/W ratio PI 4.9–7.8, PII 4.9–8.0, PIII 6.3–11.5. PI with transverse comb of about three macrochaetae laterally on the precoxa. Coxa with some scales and with strong macrochaetae in two rows along the external margin, a stout macrochaeta and some long fine setae on the inner margin subapically and group of about four or five stout curved macrochaetae at the apex over the articulation. Trochanter and femur lacking scales, femur with several strong macrochaetae ventrally and dorsally three smaller setae subdistally and one about ⅔ of the way along the margin, in addition to the fine setae over the surfaces. All tibia with four or five strong macrochaetae ventrally and another stout macrochaeta on face subapically; tibia of PI and PII with three stout macrochaetae near the outer margin. Tibia of PIII ( Fig. 107 ) with only two macrochaetae near the dorsal margin (more distal macrochaeta absent), as well as a long thinner, laterally projecting apically bifurcate macrochaeta, which is about 1.2 times longer than the tibia is wide (longer on the juvenile K377675 being almost as long as the tibia), located near the more proximal macrochaeta. Tarsus with four articles. Pretarsus with long curved lateral claws and a strong curved shorter medial claw ( Fig. 107 ).

Abdomen. Urotergites I–VII with 3+3 combs of macrochaetae as in Table 1 ( Figs 108, 109 ) noting that the macrochaeta was sometimes missing from one of the submedial combs; each comb also associated with 0–3 marginal setae, 0–5 setulae plus 1–4 cilia (e.g., Fig. 109 ). Urotergite VIII ( Fig. 110 ) with 2+2 combs, lacking the sublateral comb; urotergite IX ( Fig. 111 ) with one to five long thin infralateral setae on each side as well as a few setulae and one or two cilia. Urotergite X not very long, apically rounded, similar in both sexes ( Fig. 112 ), L/W at base about 0.6 with many strong setae along entire margin, 1+2 submarginal setae in posterolateral corners but not obviously stronger than other setae.

Urosternite I glabrous, urosternites II–VIII with 1+1 single macrochaetae ( Fig. 113 ) each associated with 0–1 marginal seta as well as a few cilia and/or setulae. Coxites of segment VIII in ♀ ( Fig. 114 ) with group of several fine setae on the rounded corners on each side of the stylus insertion. Styli in two pairs in the ♀ (VIII–IX); all styli with at least three noticeably longer and stronger setae apically ( Fig. 115 ). Styli IX almost three times as long as styli VIII and much more robust.

Coxite IX of ♀ ( Fig. 115 ), the internal process acute apically, about 3.7 times longer than the external process and 1.6–1.9 times as long as broad at its base, reaching to about 40% of the length of the stylus; external and internal margins of internal process and external margin of outer process with many moderately strong setae directed both up and down as well as a large seta adjacent to the base of the stylus, apex of outer process with several small setae. —Ovipositor long and very thin (up to 2.55 HW), surpassing the apex of stylus IX by almost twice the length of the stylus (excluding terminal macrochaetae), composed of about 38–42 divisions. Distal divisions of gonapophyses VIII and IX ( Figs 116, 117 ) with only short fine setae and setulae.

Cerci ( Fig. 95 ) with divisions from second small, several times wider than long gradually becoming longer, equally wide as long by the sixth division after which they become even longer with more annuli each with a rosette of setae and trichobothria although the large macrochaetae are restricted to the most distal annulus of each division; the most distal surviving divisions ( Fig. 118 ) with up to eleven annuli. —Medial filament of similar arrangement but subdivision occurs slightly more distad to that on the cerci.

Male. As for female except only one pair of styli (segment IX). Coxites IX ( Fig. 119 ) with acute inner process about 1.2 times longer than wide at its base and about three times longer than the more rounded external process, reaching to about 35% of the length of the stylus. Parameres small, longer than wide, with about 20 fine setae ( Fig. 120 ). Penis typical for genus with numerous glandular setae apically, each set on a protuberance.

Subadult stages. the smallest juvenile ♀ specimens examined, K377637 (HW 0.58) and K377668 (HW 0.68) have the division of urosternite VIII visible but not appearing to be complete, styli IX present but not fully developed, with no indication of styli developing on urosternite VIII and urotergite X short and semicircular. Older juvenile ♀♀ K377643 (HW 0.78), K377641 (HW 0.83), K377624 (HW 0.85), K377636 (HW 0.85) have the division of urosternite VIII appearing complete and the development of stylus VIII has usually started with one or both coxites showing a notch where the stylus will develop and the stylus is represented by a small round or triangular process with large differences in development often seen between the left and right sides of an individual; urotergite X approaching normal adult shape; in one of these specimens a feathered papilla of the maxillary palp was clearly visible. On slightly older females, K377642 (HW 0.88) and K377630 (HW 0.93) styli VIII are represented by obvious triangular processes but the ovipositor was not developed. In the specimen K377631 (HW 0.95) the developing ovipositor just surpasses the end of the internal processes of coxite IX. In females K377660 (HW 1.13) and K377657 (HW 1.13) the ovipositor is almost fully developed, surpassing the apex of styli IX by about 1.5 times the length of the stylus.

Parameres were not visible in the males K377625 (HW0.79) and K377626 (HW 0.80) but were visible in males K377629 (HW 1.00) and K377628 HW (1.04).

Etymology. The species is named for my good friends Steve and Kathy Bunton who collected on my behalf and organised the trip during which most of this material was collected.

Habitat. Heterolepisma buntonorum appears be fairly widespread in south-eastern Tasmania. Specimens were collected within cracks in the bark or under layers of bark of both living and dead Eucalyptus trees, possibly in abandoned termite galleries, and also from dry leaf litter accumulated beneath a rock overhang.

Remarks. Heterolepisma kraepelini , in the few characters adequately described (e.g., shape of ultimate article of labial palp, shape of the metasternum), appears to be similar but differs in the number of pairs of styli (three in the ♀ of H. kraepelini , two in the ♀ of H. buntonorum ) and the much longer ovipositor in H. buntonorum .

This genus now has 24 described species and will no doubt become much larger with further work. Determination of the following character states has been useful in grouping Australian specimens into species or species groups:

• presence or absence of a medial “comb” on urosternite I

• chaetotaxy of urosternites II–VII (1+1 single macrochaetae versus 1+1 combs of 2–7 macrochaetae)

• chaetotaxy of urotergite I (1+1, 2+2 or 3+3 combs)

• number of pairs of styli ( IX only in both sexes, IX in ♂ VIII – IX in ♀, VIII – IX in both sexes, VII – IX in ♀ VIII – IX in ♂, VII – IX in both sexes, VI – IX in both sexes, V – IX in both sexes, V – IX in ♀ VI – IX in ♂, IV – IX in both sexes) .

Figures

Figure 79. Heterolepisma buntonorum n.sp. Bicheno Lookout, Tasmania.

Figures 80–91.Heterolepisma buntonorum n.sp.♀ holotype unless otherwise indicated by specimen number (80) habitus; (81) macrochaeta from head above antenna; (82) scale from urotergite; (83) head; (84) antenna pedicel, scape and two intervals of flagellum; (85) idem, intervals from mid-section of flagellum; (86) idem, most distal remaining interval (near apex) showing basiconic sensory rods (sr) (K261116); (87) mandible; (88) idem, incisor and molar regions; (89) maxilla, minor setae omitted; (90) idem, lacinia; (91) labium, omitting setae from palp. Scale bars = 0.1 mm unless otherwise indicated.

Figures 92–95. Heterolepisma buntonorum n.sp. ♂ (92) macrochaetae on lateral margin of head near base of antenna; (93) chain in apical region of flagellum showing rod-like basiconic sensillae (sr) and thin-walled sensilla (tws); (94) compact sensillae of labial palp; (95) base of terminal filaments and stylus. Scale bars = 0.05 mm.

Figures 96–106. Heterolepisma buntonorum n.sp. ♀ holotype (96) pronotum with outline of limit of scales; (97) idem, trichobothrial areas; (98) mesonotum with outline of limit of scales; (99) idem, trichobothrial areas; (100) metanotum with outline of limit of scales; (101) idem, trichobothrial areas; (102) prosternum and PI; (103) idem, apex of prosternum; (104) mesosternum; (105) metasternum; (106) PIII. Scale bars = 0.1 mm.

Figures 107–114.Heterolepisma buntonorum n.sp.♀ holotype (107) PIII tibia and tarsus;(108) urotergite I; (109) urotergite VII, chaetotaxy of left side; (110) urotergite VIII; (111) urotergite IX, right infralateral chaetotaxy; (112) urotergite X; (113) urosternite VI; (114) genital region with styli, coxites VII and IX, anterior and posterior valves of ovipositor. Scale bars = 0.1 mm.

Figures 115–120. Heterolepisma buntonorum n.sp. ♀ holotype unless otherwise indicated by specimen number; (115) coxite IX of ♀ with chaetotaxy of stylus; (116) apex of anterior gonapophysis; (117) apex of posterior gonapophysis; (118) cercus, most distal surviving division; (119) coxite IX of ♂ (K261013), penis and paramere; (120) idem, apex of paramere. Scale bars = 0.1 mm.