Speciation in fractured rock landforms: towards understanding the diversity of subterranean cockroaches (Dictyoptera: Nocticolidae: Nocticola) in Western Australia Author Trotter, Andrew J. Author Main, Dean C. Author Finston, Terrie L. text Zootaxa 2017 4250 2 143 170 journal article 33308 10.11646/zootaxa.4250.2.2 50b5779a-b481-4824-a13b-666572e28edc 1175-5326 450968 36C065F7-74EC-428A-BD6C-01D578F9A27D Nocticola currani n. sp. ( Figs. 5 , 9–11 ) Type material. AUSTRALIA : Western Australia : holotype male, Chichester Range , drill hole BH0041 ( 22°30`35.70"S , 120°02`00.50"E ), 24 April 2013 ( M. Curran , D. Main ) ( WAM 84172) . Paratypes : 1 female , same data ( WAM 84171) ; 1 male , 3 nymphs, Chichester Range , drill hole BH0022 ( 22°31`48.30"S , 120°02`44.40"E ), 24 April 2013 ( M. Curran , D. Main ) ( WAM 84170) ; 1 male , 12 nymphs, Chichester Range , drill hole ML0710 ( 22°32`45.60"S , 120°01`36.70"E ), 23 April 2013 ( M. Curran , D. Main ) ( WAM 84176) ; 1 male same data ( WAM 84175) ; 1 male , 1 female , 6 nymphs, Chichester Range , drill hole ML0671 ( 22°31`21.10"S , 120°01`49.10"E ), 24 April 2013 ( M. Curran , D. Main ) ( WAM 84174) ; 1 male same data ( WAM 84173) . Etymology . The specific epithet is a patronym in honour of Michael Curran, who collected the holotype . Description of male ( holotype ). Body : Slender, approximately 3.3 times as long as wide ( Fig. 9A, B ). Colour : Pale yellowish brown. Head : Vertex of head slightly exposed, spinules present ( Fig. 9A ). Eyes reduced. Eyes are tear shaped, holotype has 15 ommatidia, located dorsal to the antennal socket (11–25 other males). Ocelli are absent. Antennae longer than body, broken on all specimens, with up to 50 segments recorded. Pronotum : Rounded, flattened towards margins, convex towards the midline, posterior margin with slight concavity ( Fig. 9A–C ), 0.81 mm long, 0.801–0.897 mm (other males), 0.956 mm wide, 0.921–1.029 mm (other males), covered in scattered small spinules with stouter longer spinules along lateral margins. Tegmina : Displays male dimorphism in tegmina. Coriaceous, reduced in length (males with tergal glands), reaching second abdominal tergum, 0.961 mm long ( Fig. 9B ), 0.914–1.163 mm (other males), 0.404 mm wide, 0.353–0.419 mm (other males) 3–4 indistinct veins ( holotype ) are present smooth, marginal setae absent ( Fig. 9B ). Males with full length tegmina are without specialised tergal glands. Tegmina hyaline, very long, extending past the body to the tip of the cerci. Eight veins are present covered in recumbent setae, marginal setae fringe short, same length as recumbent setae ( Fig. 9A ). Hind wings are present but reduced. Legs : Hind leg 1.4 times longer than total body length ( paratype ). Femora not uniformly slender, narrowing distally, anterior ventral margin of front femur with a row of minute piliform spinules and one large apical spine ( Type C1) ( Fig. 10A ). Mid and hind femurs with long setae on the ventral margins and a large spine on anterior lateral edge. Mid leg tibia with no accessory spines ( Fig. 10B ). Hind leg tibia has one medial accessory spine ( Fig. 10C ). All legs with tibia, and tarsi covered with fine hairs. Very small spinules cover tarsi on all legs. Pulvilli and arolia absent, tarsal claws simple, symmetrical, small. Front femur 0.942 mm , 0.896–1.139 mm (other males), front tibia 0.67 mm , 0.567–0.871 mm (other males), front tarsus 0.395 mm , 0.365–0.541 mm (other males), front metatarsus 0.388 mm , 0.362–0.497 mm (other males). Total front leg length 2.395 mm , 2.203–3.042 mm (other males). Mid femur 1.014 mm , 0.991–1.242 mm (other males), mid tibia 0.832 mm , 0.764–1.019 mm (other males), mid tarsus 0.49 mm , 0.466–0.694 mm (other males), mid metatarsus 0.365 mm , 0.417–0.464 mm (other males). Total mid leg length 2.701 mm , 2.656–3.419 mm (other males). Hind legs missing in holotype , hind femurs 1.13– 1.474 mm (other males), hind tibia 1.014–1.438 mm (other males), hind tarsus 0.629–0.814 mm (other males), hind metatarsus 0.464–0.622 mm (other males). Total hind leg length 3.237–4.348 mm (other males). Abdominal terga : Displays male dimorphism in abdominal terga. Unspecialized in specimens with full length tegmina ( Fig 9A ). Terga 2, 3 and 4 highly modified on short-winged morphotype ( Fig 9B ). Tergum 2 deeply concave with row of spinules along the edge of the concavity. Tergum 3 deeply concave mostly covered by segment 2 only visible at lateral edges. Tergum 4 with deep ovoid glandular pit, posterior margin of pit with a large invaginated flap lined with setae; anterior pit with a dark bow shaped gland with three raised longitudinal arms radiating from it. Cerci : Long slender, consisting of nine segments with setae of varying lengths some long and trailing. Cerci broken on holotype . Other males cerci 0.774–0.85 mm . FIGURE 9A–E. Nocticola currani n. sp. A, C, paratype (WAM ML710); B, paratype (WAM BH0022); D, E, paratype (WAM ML0671). A, habitus, dorsal view; B, habitus, dorsal view; C, head, lateral view; D subgenital plate, ventral view; E, supraanal plate, dorsal view. Scale 0.5 mm. FIGURE 10A–C. Nocticola currani n. sp. A–C, paratype (WAM ML710). A, first leg, anterior view; B, second leg, anterior view; C, third leg, anterior view. Scale 0.5 mm. FIGURE 11A–F. Nocticola currani n. sp . A–F, paratype (WAM ML0671). A, whole genitalia, dorsal view; B, R1, dorsal view; C, L2d, dorsal view; D, L3d, dorsal view; E, p, dorsal view; F, vp, dorsal view. Scale 0.2 mm. Genitalia : R 3v thin curved, lightly sclerotized, covered in fine scale-like tubercles ( Fig 11A ). R3d is positioned dorsally and is connected to the apical part of R 3v. Thin curved, lightly sclerotized, covered in fine scale-like tubercles ( Fig. 11A ). R2 located between R 3v and R3d when viewed caudally, lightly sclerotized, rounded, covered in small scale-like tubercles ( Fig. 11A ). R1 large, membranous, caudal edge heavily sclerotized with scattered setae and small tubercles towards distal end ( Fig. 11A, B ). L1 is membranous and difficult to visualise. L2d is long and hook-like, heavily sclerotized without subapical incision, with 11 long strong setae on distal end below curve of the hook. Dorsal and ventral setae scattered, setae along stem ( Fig. 11C ). L3d appears as an accessory hook-like sclerite, heavily sclerotized, spear shaped, twisted with distal third with longitudinal ribbing ( Fig. 11D ). L 2v lightly sclerotized, bow shaped ( Fig. 11A ). L 3v heavily sclerotized, harp shaped, inner edge with boot shaped extension ( Fig. 11A ). p long, lightly sclerotized, flattened with outer edge expanded. Tip long, tapering, slightly expanded, covered with long setae ( Fig 11A, E ). vp heavily sclerotized, bilobed, with dorsal and ventral ridges, some small tubercles dorsally ( Fig. 11A ). Supraanal plate : Supraanal plate in dorsal view symmetrical and trapeziform, hind margin produced and bilobed, with scattered setae dorsally and marginally. Right and left paraprocts similar, poorly defined without processes ( Fig. 9E ). Subgenital plate : symmetrical, hind margin with deeply excavate concavity, with scattered setae dorsally and marginally, styles absent ( Fig. 9D ). Description of opposite sex ( paratype ). Body : Apterous. Similar to male, but larger ( Fig. 5C ). Medial length 3.678 mm , 3.117–3.918 mm (other females). Abdomen width 1.602 mm , 1.203–1.472 (other females). Head : Eyes reduced to three separate ommatidia. Antennae longer than body, with more than 42 segments. Pronotum : Length 1.01 mm , 0.878–1.057 (other females), 1.165 mm wide, 0.985–1.276 (other females). Legs : Front leg measurements from paratype ; front femur 0.906 mm , front tibia 0.626 mm , front tarsus 0.294 mm , front metatarsus 0.377 mm . Total length front leg 2.203 mm . Mid femur 1.221 mm , 0.956–1.365 mm (other females), mid tibia 0.996 mm , 0.752–1.12 mm (other females), mid tarsus 0.59 mm , 0.518–0.712 mm (other females), mid metatarsus 0.504 mm , 0.355–0.549 mm (other females). Total mid leg length 3.311 mm , 2.581– 3.746 mm (other females). Hind leg measurements from paratype ; hind femur 1.103 mm , hind tibia 0.913 mm , hind tarsus 0.635 mm , hind metatarsus 0.499 mm . Total length hind leg 3.15 mm . Cerci : All cerci broken on female specimens. Supraanal plate: Hind margin convexly rounded with small indent medially ( Fig. 5F ). Remarks. N . currani n. sp. belongs to the uenoi -species group due to the presence of a tergal gland on terga 4 in the males ( Roth 1988 ). Tergal glands in male cockroaches produce pheromones used to attract females ( Roth 1969 ). The epigean N . australiensis is the only other Australian species of Nocticola in this group ( Roth 1988 ). Males apparently display dimorphism in tegmina length and presence of the tergal gland. The two characters are mutually exclusive in mature males; hence we do not believe they represent two life stages. Males with full length tegmina do not have tergal glands and males with tergal glands have reduced tegmina, reaching the second abdominal tergum. Specimens of each morph collected from the one bore varied in COXI sequence divergence by only 0.6% indicating they are conspecific ( Table 2 ). We are not aware of this phenomenon in any other cockroach and it may represent a case of male dimorphism. It is currently unknown which factors/variables trigger this dimorphism. TABLE 2. Pair-wise genetic distances (uncorrected p-distances; below diagonal) and standard errors (above diagonal) between haplotypes in Clade 3. The distance between haplotypes of the two morphological types of N. currani n. sp. is highlighted in yellow.

Specimen	EP6	EP10	EP7	EP8	EP9	EP11	EP12	AE8	EP13
EP6 N . currani ML0710 (CIIIa)	0.002	0.004	0.008	0.008	0.011	0.012	0.014	0.012
EP10 N . currani BH0022 (CIIIa)	0.003	0.003	0.008	0.008	0.011	0.012	0.014	0.012
EP7 N . currani ML0710 (CIIIa)	0.006	0.006	0.008	0.008	0.011	0.012	0.014	0.011
EP8 N . currani BH0041 (CIIIa)	0.039	0.039	0.039	0.005	0.011	0.011	0.013	0.012
EP9 N . currani ML0671 (CIIIa)	0.037	0.037	0.037	0.015	0.011	0.011	0.013	0.013
EP11 Nocticola sp. BH0073 (CIIIc)	0.094	0.094	0.092	0.094	0.092	0.005	0.013	0.010
EP12 Nocticola sp. ML280 (CIIIc)	0.092	0.092	0.091	0.096	0.094	0.019	0.012	0.009
AE8 Nocticola sp. BDO291 (CIIIb)	0.115	0.115	0.115	0.113	0.115	0.113	0.110	0.014
EP13 Nocticola sp. MLW19 (CIIId)	0.105	0.105	0.100	0.109	0.115	0.063	0.060	0.120

FIGURE 12. Bayesian analysis of COXI haplotypes (10 6 gens, split =0.007). Scale bar = number of substitutions per site; posterior probabilities are shown on branches. Type species are highlighted in yellow. Brackets define distinct geographic lineages within each clade as defined by geographic location. Clades are labelled with Roman numerals I–III and lineages are labeled alphabetically (a–d). Blue or red boxes indicate well-supported species containing more than one haplotype as determined by bPTP or GMYC analysis, respectively; blue or red asterisks indicate well-supported singletons as determined by bPTP or GMYC analysis, respectively; unlabeled haplotypes were not well-supported. FIGURE 13. Map of the Pilbara region in Western Australia, where Nocticola clades and lineages were collected. TABLE 3. Mean genetic distances (uncorrected p-distances; below diagonal) and standard errors (above diagonal) between lineages within each of the three major clades. Distances between lineages within each clade are highlighted in yellow. Distances between the three type specimens are highlighted in blue. Clade I Clade II Clade III