Heterolepisma cooloola, Smith & Mitchell & Lee & Espinasa, 2019
Smith, Graeme B., Mitchell, Andrew, Lee, Timothy R. C. & Espinasa, Luis, 2019, DNA Barcoding and Integrative Taxonomy of the Heterolepisma sclerophylla species complex (Zygentoma: Lepismatidae: Heterolepismatinae) and the Description of Two New Species, Records of the Australian Museum 71 (1), pp. 1-32 : 22-29
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Heterolepisma cooloola sp. nov.
Holotype. ♀ ( HW 1.20 ) ( QM 207011 on two slides) QLD: Cooloola, Freshwater track rainforest patch, 25.9492°S 153.0927°E 71 m asl, 7.vii.2013, Graeme Smith GoogleMaps . Paratypes: 14♀♀, 14♂♂, 21 subadult specimens including 1♂ ( HW 1.08 ) ( QM 207012 on two slides) same data as holotype GoogleMaps ; 1♂ ( HW 1.06 ) ( AMS K.261176, K.261177 on two slides) same data as holotype GoogleMaps ; 1 juvenile ( HW 0.74) ( AMS K.261178, K.261179 on two slides) same data as holotype GoogleMaps ; 1♀ ( HW 1.15 ) ( AMS K.261186, K.261187 on two slides) same data as holotype GoogleMaps ; 1♀ ( HW 1.09 ) ( AMS K. 377742 in ethanol) same data as holotype GoogleMaps ; 1♂ ( HW 1.20 ) ( AMS K. 377739 in ethanol) same data as holotype GoogleMaps ; 5♂♂, 5♀♀, 4 juvenile ♀♀, 3 juveniles ( AMS K.377744 all together in ethanol) same data as holotype GoogleMaps ; 1♀ ( HW1.05 ) ( AMS K.261126, K.261127 on two slides) same locality as holotype GoogleMaps , 27.i.2016, Graeme Smith; 1♂ ( HW 0.98) ( AMS K.261128, K.261129 on two slides) same data as previous GoogleMaps ; 1♀ ( HW 1.23 ) ( AMS K.261130, K.261131 on two slides) same data as previous GoogleMaps ; 1♀ ( HW 1.08 ) (gbs004893 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 1.05 ) ( AMS K.261132, K.261133 on two slides) same data as previous GoogleMaps ; 1♀ ( HW 0.93) ( AMS K.261134, K.261135 on two slides) same data as previous GoogleMaps ; 1♂ ( HW 0.85) (gbs004896 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 1.01 ) (gbs004897 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 0.85) ( AMS K. 377754 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 0.95) (gbs004899 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 0.95) (gbs004900 in 100% ethanol) same data as previous GoogleMaps ; 1 subadult ♀ ( HW 0.90) (gbs004901 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 0.95) (gbs004902 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 0.88) (gbs004903 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♀ ( HW 0.83) (gbs004904 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.75) (gbs004905 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♀ ( HW 0.75) (gbs004906 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.70) (gbs004907 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.70) (gbs004908 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♀ ( HW 0.83) (gbs004909 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.68) (gbs004910 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.75) (gbs004911 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♀ ( HW 0.68) (gbs004912 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.70) (gbs004913 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.58) (gbs004914 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ( HW 0.48) (gbs004915 in 100% ethanol) same data as previous GoogleMaps .
Other material examined. 1♂ ( HW 0.95) ( AMS K. 377743 in ethanol) QLD: Cooloola, near start of Freshwater track, 25.9439°S 153.0816°E 45 m asl, 7.vii. 2013, Graeme Smith GoogleMaps ; 1♀ ( HW 0.98) ( AMS K. 377746 in ethanol) same data as previous GoogleMaps ; 1♂ ( HW 1.01 ) ( AMS K. 377747 in ethanol) same data as previous (in ethanol) GoogleMaps ; 1♂ ( HW 1.01 ) ( AMS K. 377341 in ethanol) QLD: Carlo Point, 25.8975°S 153.0620°E 22 m asl, 6.vii.2013, Graeme Smith GoogleMaps ; 1♂ ( HW 1.03 ) ( AMS K. 377728 in ethanol) same data as previous GoogleMaps ; 3♀♀ ( HW 1.11 , 0.98 and 0.69) ( AMS K.377740 all together in ethanol) same data as previous GoogleMaps ; 1♀ ( HW1.15 ) ( AMS K. 377748 in ethanol) QLD: Cooloola, 25.9960°S 153.0716°E 63 m asl, 7.vii. 2013, Graeme Smith GoogleMaps ; 1♂ ( HW 1.03 ) ( AMS K.261188, K.261189 on two slides) same data as previous GoogleMaps ; 24 specimens ( AMS K.377745 all together in ethanol) same data as previous GoogleMaps ; 1♀ ( HW 1.15 ) ( AMS K.261180, K.261181 on two slides) QLD: Carlo Point, 25.8991°S 153.0615°E near sea level, 27.i.2016, Graeme Smith GoogleMaps ; 1♀ ( HW 1.10 ) (K.261136, K.261137 on two slides) same data as previous GoogleMaps ; 1♂ ( HW 0.95) (gbs004869 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 1.10 ) (gbs004870 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 1.13 ) (gbs004871 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 0.98) (gbs004872 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 1.05 ) ( AMS K.261138, K.261139 on two slides) same data as previous GoogleMaps ; 1♂ ( HW 0.98) (gbs004874 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 1.00) (gbs004875 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 0.90) (gbs004876 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 0.95) ( AMS K.261140, K.261141 on two slides) same data as previous GoogleMaps ; 1♀ ( HW 0.83) (gbs004878 in 100% ethanol) same data as previous GoogleMaps ; 1♀ ( HW 0.93) ( AMS K.261182, K.261183 on two slides) same data as previous GoogleMaps ; 1 juvenile ♀ ( HW 0.88) (gbs004880 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ♂ ( HW 0.83) (gbs004881 in 100% ethanol) same data as previous GoogleMaps ; 1 juvenile ( HW 0.75) (gbs004882 in 100% ethanol) same data as previous GoogleMaps ; 1♂ ( HW 0.90) ( AMS K. 377753 in 100% ethanol) QLD: Carlo Point, 25.8991°S 153.0616°E near sea level, 27.i.2016, Graeme Smith GoogleMaps .
Diagnosis. This species differs from other described species of Heterolepisma that also have 2+2 combs on urotergite I and three pairs of styli in the female and only two in the male, by the presence of lanceolate scales on the femora, tibiae and clypeus, the straight anterior margin of the head devoid of macrochaetae, the single macrochaeta mediad of the anterior trichobothrium on the pronotum, the mesosternum is also slightly different with a wider glabrous apex and the combs more compact, the posterior margin of the metasternum is rounded with a comparatively wide glabrous gap with small 1+1 combs of two to three macrochaetae. The parabolic shape of urotergite X is narrower and the terminal filaments evenly pigmented.
Appearance: Medium to large silverfish, scale covering in life uniform or slightly mottled grey with brown antennae, terminal filaments slightly darker than antennae with only a small portion of each annulus bearing the larger macrochaetae lighter in colour ( Fig. 37 View Figure 37 ).
Body length: H+B up to 9.9 mm (♀) 8.25 mm (♂); maximum HW 1.20 mm; thorax: length up to 2.85 mm (or 0.26–0.32 H+B); width up to 1.93 mm, usually slightly widest at the mesonotum; antennae damaged in all specimens, maximum preserved length of antenna 5.6 mm (or 0.68 H+B); terminal filaments damaged in all specimens, maximum preserved length of cercus 3.6 mm (or 0.48 H+B); maximum preserved length of median dorsal appendage 5.00 mm (or 0.60 H+B). Body neither elongate nor broad ( Fig. 38 View Figures 38–48 ) 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 light chestnut-brown in alcohol preserved specimens, stronger around peri-antennal and supra-ocular lines of macrochaetae and along the band of setae on the clypeus (especially laterally); pedicel and scape very lightly pigmented distally, rest of flagellum uniformly lightly pigmented becoming somewhat darker distally; all articles of maxillary palp with pigment except the most distal article, densest on article three especially distally; labium with lines of pigment around macrochaetae across the mentum, present on distal three articles of labial palp being stronger on the edges, pigment of ultimate article mostly in basal half but with a noticeable line above the more distal row of papillae. Nota with some pigment anteriorly and along margins. Legs with pigmentation along outer edge of precoxa of PI, along the length of the outer margin among the macrochaetae and only very faintly along the inner margin, distally; trochanter with light patch on margin distally; femur pigmented, darkest distally along dorsal margin and around bulge on ventral margin; tibia pigmented along edges being a little darker distally; first tarsal article pigmented distally. Urotergite X with very faint pigment along anterior lateral margins. Styli IX slightly pigmented in distal three quarters; other styli with very little or no pigment. Ovipositor with yellowish hue. Terminal filaments lightly pigmented basally becoming a lot darker distally. Some individuals show greater or lesser levels of pigmentation, with less pigmentation in juvenile specimens.
Macrochaetae: Bifid apically, or simple, hyaline, light to darker brown.
Scales: Unevenly rounded or ovoid, with numerous parallel dark brown ribs, that do not extend beyond the margin ( Fig. 39 View Figures 38–48 ); in alcohol dorsal scales and the more lateral scales of the urosternites with dark brown ribs; ventrally mostly hyaline but those towards the lateral margins with light brown ribs. Lanceolate scales present on clypeus, femora and tibia. Scales absent from flagellum of antennae, mouthparts and terminal filaments.
Head: Wider than long ( Fig. 40 View Figures 38–48 ) with marginal rows about two macrochaetae wide along the sides of the vertex, but without macrochaetae along the anterior margin, the lateral rows extending back along the margin to the eyes and extending as a single short row above the eyes, as well as a small peri-antennal group isolated from the marginal. Clypeus with numerous setae, some long and thin, others more robust but not forming combs; with a few lanceolate scales scattered among the setae. Labrum with thin setae only. Scales on top of head, those along the anterior margin overhanging the margin. Eyes dark, composed of 12 ommatidia.—Antennae long, about ⅔ H+B, scape with a subdistal rosette of setae, very conspicuous from above ( Fig. 41 View Figures 38–48 ), and numerous setae along the sides and over the ventral face ( Fig. 42 View Figures 38–48 ); pedicel short, 0.42 times the length of the scape (range 0.33–0.53), with many setae mostly distally and on the ventral face; the most apical annulus of each interval in the distal end of the flagellum with a few small inconspicuous rod-like basiconic sensilla (type B of Adel, 1984) ( Fig. 43 View Figures 38–48 ).—Mandibles ( Figs 44, 45 View Figures 38–48 ) typical for genus with well-developed molar and incisor areas; a group of about nine strong setae distally adjacent to the pectinate molar area and a bush of 50+ setae and macrochaetae externally.—Maxilla ( Figs 46–48 View Figures 38–48 ) 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 eight simple setae, the galea slightly longer than the lacinia with setulae on the outer face. Palp with rosettes of somewhat stronger setae subapically on the two basal articles, all articles with numerous fine setae, apical article of maxillary palp short, being only 0.22 times HW (range 0.20–0.25) and 4.5 times longer than wide (range 3.6–6.0) and 1.3 times longer than penultimate article (range 1.2–1.6), the ultimate article in both sexes with three “branched” papillae, those in the female much less robust and with fewer “arms” than those in the male.—Labium ( Fig. 49 View Figures 49–59 ) short and broad with rows of strong setae on the prementum and submentum, glossae and paraglossae quite broad with short curved setulae; labial palp short, apical article eccentric suboval, 1.05 times as long as wide (range L/W 0.9–1.2) with 2+3 papillae of compact type ( Fig. 50 View Figures 49–59 ) in a “cluster formation” where the slightly larger distal papillae curve around the two smaller proximal papillae and at least one curved club-like thin-walled basiconic sensillum (N.B. one palp of holotype does not show usual shape, possibly as a result of damage in the previous instar).
Thorax: Pronotum ( Fig. 51 View Figures 49–59 ) with narrow setal collar of short, apically bifurcated setae and cilia, quite weak in the medial part of the margin; lateral margins ( Fig. 52 View Figures 49–59 ) also with numerous small to medium sized, apically bifurcate setae as well as several larger more erect submarginal macrochaetae; trichobothrial areas open and in contact with the lateral margins, the anterior one ( Fig. 53 View Figures 49–59 ) located near the mid-point along the margin, almost always with one large macrochaeta located mediad of the trichobothrium (this macrochaeta missing on the left side of the holotype) and with a few cilia and setulae; posterior trichobothrial area ( Fig. 54 View Figures 49–59 ) near posterior lateral corner with a submarginal macrochaeta between the trichobothrium and the margin as well as a few setulae and cilia; posterior margin slightly concave with 1+1 combs each of one macrochaeta with a smaller seta mediad and posterior to it, the insertion of this setae smaller than that of the macrochaeta on most specimens (the smaller insertion missing from the left side of the holotype), the size of the smaller insertion becomes increasingly smaller on posterior segments such that it only appears as a very small submarginal seta on urosternite VIII; these posterior notal combs are associated with two cilia and some setulae.—Mesonotum with lateral chaetotaxy similar to pronotum ( Fig. 55 View Figures 49–59 ), the posterior trichobothrial area ( Fig. 56 View Figures 49–59 ) in the posterolateral corners with a large macrochaeta between the trichobothrium and the margins as well as some marginal and submarginal setae, cilia and setulae; the anterior trichobothrial area ( Fig. 57 View Figures 49–59 ) about ¾ the distance posteriorly along the margin, with a submarginal macrochaeta (m-1) between it and the margin, also with a few setulae and cilia; anterior to this trichobothrial area are two combs (m-2, m-3) each of two macrochaetae, a further two or three submarginal macrochaetae more anterior along the margin; posterior combs as for pronotum ( Fig. 58 View Figures 49–59 ).— Metanotum ( Fig. 59 View Figures 49–59 ) similar to mesonotum (the comb at position m-2 on the left side of the holotype composed of only one macrochaeta, suggesting that a degree of variation exists within the species).
Presternum narrow, with transverse row of strong setae and numerous cilia.—All thoracic sterna with hyaline scales. Prothoracic sternum pointed cordiform, only slightly longer than wide at its base (L/W 1.08 range 1.4–1.15) and reaching almost to the end of the coxa, rounded apically and with a medial furrow ( Fig. 60 View Figures 60–65 ), most of lateral margins with numerous small marginal setae and cilia, with 4–6 larger submarginal macrochaetae forming weak combs parallel to the edges in the distal quarter.—Mesosternum ( Fig. 61 View Figures 60–65 ) slightly longer than broad (1.09 range 0.98–1.18) with a truncate or evenly slightly concave posterior margin, with 1+1 distal combs of two to three submarginal macrochaeta associated with some marginal setae and cilia ( Fig. 62 View Figures 60–65 ).— Metasternum ( Figs 63, 64 View Figures 60–65 ) wider than long (L/W 0.75 range 0.69–0.84) but otherwise similar to the mesosternum; the gap between the combs 7.3 times the average width of each comb (range 5.4–11.1).
Legs fairly long ( Figs 60, 61, 63 View Figures 60–65 ), tibia L/W ratio of legs PI 2.6 (range 2.5–3.0), PII 3.0 (range 2.4–3.5, PIII 3.6 (range 3.1–4.1); tarsi L/W ratio PI 6.7 (range 5.4–7.6), PII 6.7 (range 6.0–8.0), PIII 8.8 (range 7.4–10.0). Legs increasingly longer from front to back, mean ratio PI/PIII (tibia 0.58, tarsus 0.68). PI with transverse comb of about six macrochaetae laterally on the precoxa. Coxa of all legs covered with hyaline scales and with strong macrochaetae and numerous cilia in a row about two macrochaetae wide along the external margin, a stout macrochaeta and some long fine setae on the inner margin subapically and group of about four to six stout curved macrochaetae at the apex over the articulation. Trochanter lacking scales, with fine and one stronger seta over the surface. Femur with numerous lanceolate scales on the anterior half and along the margin, rest of surface with fine setae; anterior distal end with three to six strong, quite deeply bifurcate stout macrochaetae as well as some strong setae; posterior margin with several strong macrochaetae as illustrated. Tibia with numerous setae and lanceolate scales over the ventral surface, with two stout macrochaetae on or near the anterior margin and three or four stout macrochaetae along the posterior margin (some paired with thinner macrochaetae on the dorsal side of the margin; apical spur with several setae. Tibia of PIII with a long thin, laterally projecting trichobothria-like seta inserted dorsal to the proximal stout macrochaeta on the anterior margin, which is more than twice as long as the tibia is wide. Tarsus with four articles, all with numerous setae (without lanceolate scales). Pretarsus with long curved lateral claws and a strong curved shorter medial claw ( Fig. 65 View Figures 60–65 ).
Abdomen: Urotergite I usually with 2+2 combs of 1–2 macrochaetae (sublateral missing on right side of holotype), urotergites II–VII with 3+3 combs of macrochaetae ( Fig. 66 View Figures 66–77 ) as in Table 8, 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., Figs 67–69 View Figures 66–77 ). Urotergite VIII ( Fig. 70 View Figures 66–77 ) with 2+2 combs, lacking the sublateral comb, each comb associated with 0–2 marginal setae, 2–5 setulae and 2–4 cilia; urotergite IX with two long thin infralateral setae on each side as well as a 1–2 cilia ( Fig. 71 View Figures 66–77 ). Urotergite X fairly long and slender parabolic in both sexes ( Figs 72, 73 View Figures 66–77 ), L/W at base about 0.6 (range 0.54–0.70) with many strong setae along entire margin, often without obvious strong submarginal macrochaetae in the postero-lateral corners but sometimes with up to two submarginal insertions visible on each side.
Urosternite I glabrous, urosternites II–VIII ( Fig. 74 View Figures 66–77 ) with 1+1 single macrochaetae ( Fig. 75 View Figures 66–77 ) (although missing from left side of urosternite II on holotype), each associated with 0–1 marginal seta as well as a few cilia and/or setulae. Coxites of segment VII, VIII and IX in ♀ ( Fig. 76 View Figures 66–77 ) with group of several fine setae on the rounded corners on each side of the stylus insertion ( Fig. 77 View Figures 66–77 ). Styli in three pairs in the ♀ ( VII–IX); all styli with several noticeably longer and stronger setae apically. Styli IX three times as long as styli VII (range 2.3–3.7) and about two and a half times as long as stylus VIII (range 2.0–2.9) and much more robust ( Fig. 78 View Figures 78–85 ) .
Coxite IX of ♀ ( Fig. 76 View Figures 66–77 ), the internal process acute apically, about 4.2 times longer than the external process (range 3.3–6.0) and 1.8 times as long as broad at its base (range 1.6–2.1), reaching almost to half the length of the stylus; external and internal margins of internal process and external margin and apex of outer process with many moderately strong setae directed both up and down.— Ovipositor ( Fig. 76 View Figures 66–77 ) very long and thin (up to 2.30 HW), surpassing the apex of stylus IX by at least the length of the stylus (excluding terminal macrochaetae), composed of about 34–40 divisions. Distal divisions of gonapophyses VIII and IX ( Figs 79, 80 View Figures 78–85 ) with only short fine setae and setulae.
Cerci ( Figs 81, 82 View Figures 78–85 ) with basal divisions shorter than long, gradually becoming longer distally, equally wide as long by about the sixth division after which they become even longer with more annuli each with a rosette of setae and some with trichobothria with the large macrochaetae restricted to the most distal annulus of each division; the most distal surviving divisions with up to eight annuli.—Medial filament of similar arrangement ( Figs 81, 83 View Figures 78–85 ).
Male: As for female except only two pair of styli (segments VII and IX). Coxites IX ( Fig. 84 View Figures 78–85 ) with acute inner process about 1.8 times longer than wide at its base (range 1.70–1.93) and about four times longer than the external process which has a small preapical constriction, reaching to about half the length of the stylus. Both processes with several strong setae mostly apically emerging from both the dorsal and ventral surfaces of the processes close to or on the margin. Parameres small, slightly longer than wide, with about eight fine setae ( Fig. 85 View Figures 78–85 ). Penis typical for genus with numerous glandular setae apically, each set on a protuberance ( Fig. 84 View Figures 78–85 ).
Subadult stages: Very small specimens (HW 0.68) only have styli on coxites IX; the coxites of segment VIII are already clearly divided in a juvenile ♀ of HW 0.53. The ovipositor is just beginning to appear in a ♀ of HW 0.83 and styli VIII are present but not styli VII; a ♀ with HW of 0.88 had an ovipositor that just attained the end of styli IX but still lacked styli VII; by HW 0.93 all styli were present and the ovipositor was much longer than the end of stylus IX. In a ♂ with HW 0.96 one stylus VIII was developed but the other was only represented by a small triangular appendage. Presumably specimens of both sexes could be considered as sexually mature once HW is greater than 0.93–0.96 mm or once all styli are clearly developed.
Habitat. Heterolepisma cooloola was fairly common in the Rainbow Beach area with specimens collected in dry leaf litter accumulating in places largely protected from rainfall such as within burned out ground level tree hollows. It was also taken from the bark of ti-trees, Casuarina and Eucalyptus in heathland and on the edge of rainforest using pyrethrum sprays.
Etymology. The species name is derived from the proper noun Cooloola referring the locality in which it was collected. Comments. Heterolepisma cooloola is in many ways similar to H. sclerophylla (small triangular prothoracic sternum, glabrous urosternite I, urosternites II–VIII with 1+1 macrochaetae, three pairs of styli in the female and two in the male) but in other aspects it shares characters with H. parva Smith from Barrow Island (notably the glabrous anterior margin to the frons, a macrochaeta mediad of the anterior trichobothrium on the pronotum and the presence of lanceolate scales). Heterolepisma parva is however, one of the species with a medial comb on urosternite I and 1+1 combs of several macrochaetae on urosternites II–VII (VIII), a group that Mendes (pers. comm.) has suggested may be a separate group within Heterolepisma . The combined molecular and morphological data support the view that the presence of lanceolate scales and the absence of macrochaetae from the anterior margin of the frons are more significant to phylogeny than the arrangement of styli and the shape of the thoracic sternites in Heterolepisma .
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