Moore (1963) diagnosed Mecyclothorax based on: 1, apical palpomeres glabrous; 2, pronotum cordate; 3, male parameres narrowed apically. These characters were chosen to differentiate Mecyclothorax from Neonomius Moore, a genus now placed far from Mecyclothorax in the subtribe Moriomorphina. The first character works well for Mecyclothorax, with the rare occurrence of very short setae on the apical palpomeres, those setae about 0.10× the palpʼs maximal diameter. Since Mooreʼs pioneering exposition, pronotal shape has been shown worthless for diagnosing Mecyclothorax given the great diversity in shape among the Tahitian (Perrault 1984, Liebherr 2013), Hawaiian (Liebherr 2015), and New Caledonian species (Liebherr 2018). The male parameres differ in configuration among the various subgenera of Mecyclothorax, but as Moore proposed, both right and left parameres are elongate and setose apically (Liebherr 2018, fig. 5). Other characters that diagnose Mecyclothorax from various other taxa in tribe Moriomorphini, both symplesiomorphically and synapomorphically (Liebherr 2011a), include: 1, labrum truncate to moderately and broadly emarginate apically; 2, ligula truncate medially between the 2 marginal setae, paraglossae elongate; 3, pronotal base at least moderately punctate; 4, elytral humeral margin smooth, without tooth at humeral angle; 5, female apical ventrite with median patch of 4–5 smaller setae.

Sloane (1915) described Mecyclothorax ovalis from Manjimup, WA, with label data (ANIC): Manjimup, W.A. / 31.12.13 T. G. S. // Mecyclothorax / ovalis Sl. Type // Holotype [pink label]. The pink holotype label is consistent with those used by P. J. Darlington, Jr. during his curation of the Sloane collection in 1957 (Darlington 1962: 328). The specimen is mostly destroyed, with only partial remnants of the legs remaining glued to the card. As such, because the holotype specimen can no longer be used to diagnose the species nor the species’ generic placement, a neotype is designated. As this neotype designation is required to stabilize both the concept of this species and the species’ new generic combination, such an action is valid under Article 75 of The Code (ICZN 1999). Therefore a Neotype male (MCZ specimen deposited in ANIC) is hereby designated: Mullewa, W.A. / Sept. 19 1931 // Australia / Harvard Exp., / Darlington // MCZ // Neotype / Mecyclothorax / ovalis Sloane / det. J.K. Liebherr 2004 [black-margined red label]. These specimens, both neotype and others in the series (MCZ) exhibit densely setose apical palpomeres as well as the small ovoid body diagnostic for Neonomius Moore (1963). The male genitalia conform to the configuration observed in the subtribe Moriomorphina (Liebherr 2011a), with both parameres broad, parallel-sided with rounded apices, and the right paramere ventrally setose, as observed in the generic type species N. laevicollis (Sloane) (Moore 1963, fig. 7). Based on these characters, Sloaneʼs species is newly combined as Neonomius ovalis (Sloane), comb. n. Among species of Neonomius, N. ovalis is diagnosable from the sympatric N. australis (Sloane) by: 1, the smaller body size, standardized body length 3.2–3.6 mm for the N. ovalis neotype series – agreeing with Sloane (1915; 451) – versus 5.0 mm for N. australis (Sloane 1915: 450); and 2, the reddish-brown body color versus “black, nitid; legs piceous-red; antennae reddish (Sloane 1915, 450)” for N. australis. Neonomius ovalis can be diagnosed from the other two southeastern Australian species placed in the genus – N. laevicollis (Sloane) and N. laticollis (Sloane) – by the apically less convex elytral interval 8, versus the laterally compressed, subcarinate interval 8 characterizing those two species. The type locality for M. ovalis (Sloane) becomes Mullewa. That Moore et al. (1987) did not recognize Sloane’s Mecyclothorax ovalis as a member taxon of Neonomius Moore is based on the lack of any other specimens in the ANIC beyond the destroyed holotype of M. ovalis Sloane, whereas Darlington made this nomenclatural connection, but only for specimens he deposited in the MCZ.

All other names placed under Mecyclothorax Sharp in Moore et al. (1987: 147–149) are treated below.

Beetles assigned to this subgenus have the prosternum punctate, with punctures either: 1, distributed over the lateral reaches of the prosternum and medial reaches of the proepisternum; 2, more generally distributed across both sclerites; 3, lining a median depression anterad the prosternal process; 4, lining the prosternal-proepisternal suture; or 5, lining a transverse preapical depression (Liebherr 2018, fig. 2D–F). The pronotum may be distinctly punctate (Fig. 2A–B) or not (Fig. 2C–I), but in all species the median pronotal base is coplanar to only slightly depressed relative to the pronotal disc, not greatly depressed as in Australian species of subgenus Mecyclothorax (Fig. 2J–M). The vertex is transversely flat to convex, without a transverse dorsal impression, or neck. Generally the species conform to the full complement of standard setae; formula ++/++/+2++. However the basal pronotal seta is absent in M. darlingtoni and M. jameswalkeri, the parascutellar seta is polymorphically present or absent within M. punctatus, and M. isolatus exhibits only a single dorsal elytral seta. Body size ranges from small to moderately large, with standardized body length 2.7–6.0 mm.

Figure 2. 

Pronota, dorsal view, of mainland Australian Mecyclothorax spp.: A, M. (Eucyclothorax) moorei; B, M. (Eucyclothorax) punctatus; C, M. (Eucyclothorax) curtus; D, M. (Eucyclothorax) blackburni; E, M. (Eucyclothorax) darlingtoni; F, M. (Eucyclothorax) lophoides; G, M. (Eucyclothorax) eyrensis lectotype; H, M. (Eucyclothorax) peryphoides holotype; I, M. (Eucyclothorax) cordicollis; J, Mecyclothorax (s. s.) lateralis paralectotype; K, M. (s. s.) minutus lectotype; L, M. (s. s.) ambiguus; M, M. (s. s.) punctipennis.

All Eucyclothorax spp. are characterized by male genitalia that possess a flagellum, flagellar sheath, and dorsal plate at the apex of the internal sac, as well as robust aedeagal median lobes, i.e. dorsoventrally broad (Figs 7, 14). The female reproductive tract is configured with the spermathecal duct joined ventrally to the juncture of the common oviduct and the bursa copulatrix (Fig. 9). A helminthoid sclerite is present on the bursal wall ventrad the juncture of the spermathecal duct and common oviduct. Like all other Mecyclothorax, the gonocoxae have setae along the apicolateral margin of the basal gonocoxite (Fig. 10), and the apical gonocoxite has 1–3 (usually 2) lateral ensiform setae.

(n = 2). This species is diagnosed by the broadly punctate pronotal disc (Fig. 2A), punctate head with a transverse line of about 5 large punctures between the posterior supraorbital setae, and convex, smooth elytra with only striae 1–4 in evidence, striae 2–4 only a series of small isolated punctures. The prosternum and proepisternum are also broadly punctate, the punctures large and separated from each other by a distances equal to the punctural diameters (Liebherr 2018, fig. 2F). The punctate pronotal disc, head, and prosternum + proepisternum are shared with M. punctatus, but the pronotal punctures are more numerous and less separated in M. punctatus (Fig. 2A–B), with about 30 punctures each side in M. moorei, and about 60 each side in M. punctatus. In addition, the elytral striae are more distinct in M. punctatus, with striae 1–5 present in the basal half of the elytra, though the punctures of stria 5 are smaller and shallower. The elytra of M. moorei are more ovoid, with the lateral margins more narrowly rounded behind the subangulate humeri (Fig. 3A–B). Whereas the parascutellar seta is present or absent in M. punctatus, and if present appearing short and narrow, this seta is well developed in M. moorei, with the setal articulatory socket set within a depression coincident with the base of the parascutellar striole, the seta as long as the breadth of 2–3 elytral intervals. The eyes of M. moorei are convex but they cover only the anterior 4/5 of the ocular lobe: EyL/OLL = 0.83–0.86. Finally, males of this species, M. punctatus, and M. curtus all exhibit 2 setae each side of the apical abdominal ventrite, an unusual character state within Mecyclothorax, and evidence of their monophyletic relationship within subgenus Eucyclothorax (Fig. 1B, character 83). Standardized body length 3.2–3.3 mm. Setal formula ++/++/+2++.

Male genitalia (n = 2). Aedeagal median lobe apex narrowly rounded and only slightly projected beyond apical ostial margin (Fig. 7A); aedeagal internal sac elongate, membranous, with apical dorsal plate, flagellum, and flagellar sheath (Fig. 7B); right paramere narrow, elongate, with 6 setae along the ventral margin, 1–2 apical setae, and the dorsal margin glabrous except for a small apical seta (Fig. 8A); left paramere broadly quadrate basally, the apical 1/3 of length very narrow in contrast to broad base.

Female reproductive tract (n = 1). Bursa copulatrix elongate, columnar (Fig. 9A); helminthoid sclerite elongate; spermathecal duct basally expanded and sclerotized, the more apical portion of duct membranous and of lesser diameter; basal gonocoxite with 2 larger setae laterally along apical margin (Fig. 10A), a very small seta medially near margin; apical gonocoxite broad basally, breadth more than half length; lateral ensiform setae elongate, ~ 0.60× length of apical gonocoxite; apical nematiform setae in subbasal sensory furrow.

Holotype male (AMS): site 32AR NSW Banda Rd about 4.5 km E Hastings Forest Hwy 31°09’S 152°25’E Mount Boss State Forest 17 1100m (NPWS Survey) 4 Feb–9 Apr 1993 M. Gray, G. Cassis (AMS K241125) (Baehr 2009).

M. moorei is restricted to northeastern New South Wales (Fig. 11A), with populations allopatrically distributed relative to eastern populations of the more southerly distributed M. punctatus. Recorded elevations of collecting localities range from 110 m near Ramornie and 1100 m in Mt. Boss State Forest (Baehr 2009). Philip Darlington collected this species in Bellangry Forest NW of Wauchope (MCZ). All specimens are vestigially winged.

(n = 5). This species is most similar to M. moorei, with several diagnostic characters differentiating the two listed there. The eyes are large and convex, largely covering the ocular lobes; EyL/OLL = 0.92–0.96. The elytra are broad, with broadly rounded humeri (Fig. 3B). The single available female specimen assignable to M. p. peckorum has the elytral base broad, slightly different from the nominate form. However the pronotal and elytral strial punctures are deeper and less dense in M. p. peckorum–as reported in Baehr (2016a)–resulting in recognition of the two forms here. In addition, the parascutellar seta is much less in evidence among individuals of this species. When present it is short and fine, and it is certainly absent from the majority of examined specimens. Standardized body length 2.7–3.2 mm. Setal formula ++/++/±2++.

Figure 3. 

Mecyclothorax (Eucyclothorax) spp., dorsal view: A, M. moorei; B, M. punctatus, NSW; C, M. curtus; D, M. blackburni; E, M. isolatus holotype.

Male genitalia (n = 1). Aedeagal median lobe apex narrowly rounded and not projected beyond apical ostial margin (Fig. 7C); right paramere narrow, elongate, with ~10 setae along the ventral margin, 2 apical setae, and the dorsal margin with 3 small setae (Fig. 8B); left paramere broadly expanded basally, its dorsal margin convex, the apical 1/3 of length very narrow in contrast to broad base. Baehr (2016a, fig. 2) documents a short leftward expansion on the median lobe apex of M. p. peckorum. The parameral setation of the two subspecific forms is identical (Fig. 8B; Baehr 2016a, fig. 2).

Female reproductive tract (n = 1). Bursa copulatrix elongate, columnar (Fig. 9B); helminthoid sclerite elongate; spermathecal duct basally expanded and sclerotized, the more apical portion of duct membranous and of lesser diameter; basal gonocoxite with 3 larger setae laterally along apical margin (Fig. 10B); apical gonocoxite broad basally, breadth more than half length; lateral ensiform setae broad, elongate, length about 0.5× length of apical gonocoxite, setal surface longitudinal striate; apical nematiform setae in subbasal sensory furrow.

For M. punctatus, Lectotype female (SAMA) hereby designated: white card with “5104” and “N.S.W.” in red ink // N.S. Wales // punctatus Sloane // J. 7252 / Mecyclothorax / punctatus / M.S. Wales / Cotype [red ink] // Lectotype ♀ / Cyclothorax / punctatus Sloane / det. J.K. Liebherr 2004 [red label with black border]. For M. punctatus peckorum, Holotype male (ANIC, not seen): Pemberton Warren N.P. WA 12 Jul. 1980 S. & J. Peck SBP106 // Berlesate karri litter and moss (Baehr 2016a).

Following Baehrʼs (2016a) circumscription of this species, it is distributed across an easterly set of populations in southern New South Wales and northern Victoria (Fig. 11A), plus a disjunct set of western localities in the southwest region of Western Australia. Sloane wrote of the type series, “I have found it in considerable numbers under logs and the leaves of saplings at a place about twenty-five miles north-west from the town of Urana. It is sluggish in its movements (Sloane, 1895: 449–450).” Thus the type locality is very close to one of the localities where he collected types of M. cordicollis (Sloane 1900). More recently, specimens from the western localities have been collected via litter sifting of jarrah forest litter (Eucalyptus marginata Donn ex Sm.). All specimens are vestigially winged.

(n = 4). The transverse pronotum–MPW/PL = 1.39–1.47 with nearly impunctate median base and broadly convex margins (Fig. 2C), and broad, subparallel elytra (Fig. 3C), are unique within Mecyclothorax, unmistakably diagnosing this very rarely collected species. The pronotal hind angles are defined by denticles on a more broadly convex margin, the basal pronotal setae associated with a broadened marginal bead at their point of insertion. There are 5–6 minute punctures each side on the median base inside the laterobasal depressions, which are marked by 2–3 oblique lines of larger punctures associated with the depression (Fig. 2C, left side). Ventral prothoracic punctation is restricted to the lateral reaches of the prosternum (Liebherr 2018, fig. 2D). The lateral margins of the abdominal ventrites are also distinctly punctate, the punctures distributed densely and irregularly enough to give the surface a dented look. As in the above two species, both males and females have 4 setae along the apical margin of the apical abdominal ventrite. As in M. punctatus, the eyes are large and convex, EyL/OLL = 0.87–0.91. However unlike M. punctatus and M. moorei, the elytral striae are fully developed, with all striae evident nearly to the elytral apex. There is a well-developed carina immediately laterad stria 7 that extends from the position of the posterior series of lateral elytral setae to the elytral margin distad the well-developed elytral plica, parsimoniously interpreted as a parallelism also observed in M. blackburni (Fig. 1B, Liebherr 2018, fig. 2L). The apical palpomeres bear a sparse pelage of very short setae, a character otherwise only observed within the genus in Norfolk Islandʼs M. monteithi. Standardized body length 3.8–4.1 mm. Setal formula ++/++/+2++.

Male genitalia (n = 1). Aedeagal median lobe apex narrowly rounded with broad dorsal expansion, the apical face of the lobe concave (Fig. 7D); flagellum short, associated with very elongate sclerotized saccal surface herein interpreted as flagellar sheath (single available male with sac uneverted); right paramere elongate but broad apically (Fig. 8C), the paramere more robust than in other Eucyclothorax (Figs 8, 13), and with ventral surface bearing only a single short seta complementing the 2 long apical setae, and the dorsal surface glabrous; left paramere broad basally (as in males of M. moorei and M. punctatus), but also broad apically, the apex more than half the breadth of base, a single longer seta present at apex.

Female reproductive tract (n = 1). Bursa copulatrix elongate, columnar (Fig. 9C); helminthoid sclerite with distal projection; spermathecal duct elongate, more than twice length of spermathecal reservoir, the duct moderately sclerotized, undulated along length; basal gonocoxite with a single large seta laterally along apical margin (Fig. 10C); apical gonocoxite very broad, the base extended and recurved basally so that lateral margin is distinctly concave; lateral ensiform setae broad, dorsal ensiform seta situated medially so that its apex extends beyond medial margin; apical nematiform setae in subbasal sensory furrow.

Holotype (ANIC): specimen on white triangle, lacking head and prothorax // C. curtus Sl. (type) / Bendigo. W.W.F. / 1176; “1176” is in Sloaneʼs specimen list for 1893/94 (ANIC, unpubl. data).

The lone holotype was collected by W.W. Froggatt at Bendigo, Victoria. Besides the holotype, I have had the opportunity to examine only a second specimen from Bendigo (MVM), two specimens from Sea Lake, Victoria (Fig. 11B; ANIC, MVM) and a single female from 27 km W Manangatan (ANIC). This last specimen is labeled “South Australia”, but based on the locality data it must be from Victoria. Nothing is recorded concerning this speciesʼ habits, though occupation of a terrestrial microhabitat associated with forest vegetation near water would be consistent with the habits of its adelphotaxon, the sister-species pair M. moorei and M. punctatus (Fig. 1B). All specimens are vestigially winged.

(n = 5). Beetles of this species are very narrow-bodied, with a narrow, cordate pronotum (Fig, 2D), and narrow, subparallel elytra and an elongate head (Fig. 3D). The pronotal median base is coplanar with the disc, but distinguished by the presence of about 14 large deep punctures each side. The right pronotal hind angles protrude both laterally and posteriorly in association with a broadened marginal bead at the articulatory socket of the basal pronotal seta. Otherwise the pronotal lateral margin is extremely narrow, defined only by a marginal bead. Basally the elytral striae consist of series of closely spaced punctures, their separations equal to their diameters. Puncture size decreases, and puncture distances increase laterally and apically on the elytra, with stria 7 represented by only a few small punctures at mid-length. Interval 8 is broadly convex apically, and the elytral plica is well developed and evident in dorsal view. Ventrally, the body punctation includes a punctate median depression anterad the prosternal process, punctures along the lateral reaches of the prosternum, and a punctate anteapical groove (Liebherr 2018, fig. 2E). The lateral reaches of the mesosternum, metasternum, and all of the metepisternum are also intensely punctate. The basal 3 abdominal ventrites are covered with numerous small punctures laterally, and the suture between visible ventrites 2 and 3 is traceable only as a shallow groove. The apical margin of the male apical abdominal ventrite bears the usual 2 setae, 1 each side, but also 4 small medial setae in the position observed in Mecyclothorax females. Standardized body length 4.2–4.9 mm. Setal formula ++/++/+2++.

Male genitalia (n = 2). Aedeagal median lobe very broad and only slightly curved, the apex with subacuminate ventroapical projection and a broadly convex dorsoapical expansion resulting in a broadly concave apical face (Fig. 7E); flagellum short, flagellar sheath robust, and dorsal plate lightly sclerotized, difficult to discern in single available uneverted male; right paramere narrow, with 4 small setae ventrally and dorsal surface glabrous (Fig. 8D); left paramere broad basally, narrowly attenuate apically.

Female reproductive tract (n = 1). Bursa copulatrix elongate, extended dorsodistally beyond juncture of common oviduct and bursa (Fig. 9D); helminthoid sclerite robust, with distal projection; spermathecal duct straight and slightly longer than spermathecal reservoir, evenly, moderately sclerotized; basal gonocoxite with 1 larger seta apically, a smaller seta present or not (Fig. 10C); apical gonocoxite narrowly triangular, narrowed apically to acuminate apex; lateral ensiform setae narrow, moderately elongate; apical nematiform setae in apical sensory furrow.

Holotype male (SAMA): card mounted // TY Pinjarrah // Holotype M. blackburni / PJD Sl. [red label].

Sloane (1903) proposed the replacement of Cyclothorax MacLeay (1871) (not Frauenfeld 1868) with Sharpʼs Mecyclothorax (1903), mentioning the new combinations of M. lateralis (Castelnau), M. fortis (Blackburn) = M. minutus (Castelnau) [NEW SYNONYMY herein], M. punctatus (Sloane), and M. curtus (Sloane). By not mentioning M. blackburni, nor also M. eyrensis (Blackburn), M. peryphoides (Blackburn), M. cordicollis (Sloane), M. minutus (Castelnau), and M. punctipennis (MacLeay), Csiki (1929) became the first to propose these combinations (see also below).

M. blackburni is known only from coastal Western Australia (Fig. 11B) from Perth south to Harvey (ANIC, 2 specimens). The Perth locality is denoted as “Bridgelʼaʼ (MVM, 1 specimen), which is here interpreted to be an abbreviation of Bridgeleigh, a remnant area of bush vegetation in Swan Valley, Wanneroo. Commander J.J. Walker, Royal Navy, collected a specimen at Fremantle in 1914 (BMNH). When Sloane (1898) described the species based on one specimen received from Arthur Lea he listed no biological information, and none accompanies the other four available specimens. Those four specimens are all macropterous.

(n = 1). The larger body size, standardized body length 6.0 mm, ferruginous body color, and broadly transverse, ovoid pronotum (Fig. 3E) serve to diagnose this species from others of subgenus Eucyclothorax, except perhaps the smaller-bodied M. curtus (Fig. 3C). However the lateral margin of the pronotum is broadly rounded behind in this species, with the margin explanate in the region of the basal pronotal seta. The metathorax is remarkably abbreviated, more so than in any other Australian species of Mecyclothorax, with the metepisternum transversely broader than its lateral length. The elytra exhibit a subcarinate ridge along the eighth interval dorsad the subapical sinuation, reminiscent of such a carinate ridge observed in M. blackburni. The carinate ridge lies dorsad a well-developed internal elytral plica, with the plica and elytral margin fitting into a corresponding invagination along the margin of the apical abdominal ventrite, thereby forming, in concert with the conjoining of the elytra at the suture, a very robust, shell-like hindbody. Finally, there is a single dorsal elytral seta on each elytron (Fig. 3E), situated near midlength as observed in species of subgenus Qecyclothorax (Fig. 5D). Setal formula +/+/+/+/+/1/+/+.

Head capsule elongate, frons with medial depression but otherwise convex mesad the deep, sinuous frontal grooves; frontal grooves deeply and obliquely continued onto clypeus toward lateral clypeal margins; labrum slightly emarginate apically; antennae filiform, antennomere 9 length 2.36× breadth; mandibles moderately elongate, overall length 1.57× distance from anterior condyle to lateroapical labral margin; eyes well developed and moderately convex, ocular lobe broadly projected, outer eye surface of same curvature behind as posterior portion of lobe meeting gena, ocular ratio = 1.47, ocular lobe ratio = 0.79; mentum tooth with sides obtuse, apex broadly rounded; ligular apex moderately narrowed, 2 ligular setae separated by 3 setal diameters; paraglossae thin, extended 1/2× as far past ligular margin as distance from base to ligular margin; mentum broad, breadth/length across the lateral lobes = 3.25. Pronotum broadly ovoid, MPW/PL = 1.31, without any indication of hind angles save a slight change of curvature of the lateral margin at the hind seta; articulatory socket of lateral seta 1 setal diameter mesad deepest part of marginal depression; median base convex, unmargined medially though broadly upraised mesad laterobasal depressions; base convex anterad basal margin, slightly depressed relative to convex disc, and separated from disc by broad, smooth oblique depressions that extend to laterobasal depressions; about 8 indistinct punctures each side from midline to mesal margins of laterobasal depressions, the depressions broadly extended to explanate lateral margin, with a low upraised tubercle in the middle of each depression; median longitudinal impression fine, well indicated, crossed by transverse wrinkles on disc; anterior transverse impression broad, shallow medially, not indicated laterally; anterior callosity slightly, broadly convex, a well-defined marginal bead along front of pronotum; front angle moderately protruded, tightly rounded; prosternal process broadly depressed between procoxae; prosternum smooth and convex medially, indistinctly punctate anterolaterally with an indistinct anteapical groove consisting of broad punctures that anastomose into a groove along lateral reaches; proepisternum impunctate, however prosternal-proepisternal suture lined with about 5 indistinct punctures; proepimeron with broadly raised posterior bead, the suture with proepisternum smooth. Mesepisternum punctate at its deepest portion, about 6 deep punctures in 1–2 dorsoventral rows. Elytra with striae 1–5 composed of isolated punctures in basal half, less punctate though traceable in apical 1/3 of length, stria 6 represented by small, isolated punctures at midlength, stria 7 absent except near apex mesad subcarinate eighth interval dorsad subapical sinuation; sutural stria broadly depressed in apical half in association with convex sutural interval, the elytra conjoined apically; stria 8 a series of deep, interrupted punctures at midlength, deep and continuous mesad the posterior series of lateral elytral setae; lateral elytral setae arrayed in 7 + 6 (anterior series setae and posterior series setae), with the posterior seta of the anterior series slightly separated from the rest; subapical sinuation angulate, abruptly curved anteriorly, with well-developed internal elytral plica visible in quarter view, though obscured by the elytral margin in dorsal view. Metepisternum short, trapezoidal, maximum breadth 1.1× lateral length, metepimeron broadly convex posteriorly; metasternal process with sides acute, apex narrow, triangular with margin very broad medially in apex of process. Abdomen with broad linear depressions on lateral reaches of visible ventrites 3–6; suture between ventrites 1 and 2 deeply sinuous laterally, ventrite 2 depressed within sinuosity; female with 2 setae each side and a median patch of 4-5 smaller setae; apical margin of the female apical ventrite with deep emargination each side bordered laterally by a vertical, sclerotized border, these emarginations and lateral wall fitting into the elytral plica above. Microsculpture absent from frons, the surface glossy, micropunctures visible across the surface; pronotal disc and base with indistinct transverse microsculpture consisting of transverse lines and elongate meshes, these visible in surface irregularities such as wrinkles and depressions; elytral disc glossy with fine transverse lines faintly visible outside areas of reflection, elytral apex with transverse sculpticells visible in irregularly depressed areas associated with striae; metasternum glossy with indistinct transverse sculpticells, their breadth 2–3× length; abdominal ventrites glossy with swirling transverse mesh and transverse lines. Coloration of head rufous; antennomere 1 flavous, antennomeres 2–3 rufoflavous, 4–11 with brunneous cast; pronotal disc dark rufous, margins rufoflavous; elytral disc rufobrunneous, sutural interval concolorous, interval 9 and marginal depression, and apex narrowly rufoflavous; proepipleural margin rufous, rufoflavous ventrally, proepisternum rufous; elytral epipleuron broadly flavous, margin darker, brunneous, metepisternum rufoflavous; abdominal ventrites rufoflavous with dark rufous posterior margins, apical ventrite with apical half rufoflavous; femora flavous; tibiae brunneous.

Female reproductive tract (n = 1). The unique female holotype was not dissected, however the gonocoxae are exerted from the specimen allowing the following characters to be assessed: basal gonocoxite with 2 stout apicolateral setae, medioapical surface glabrous (as in Fig. 10A); apical gonocoxite extended laterally at base, basal width about 0.7× length, 2 stout lateral ensiform setae along lateral margin of coxite and an elongate dorsal ensiform seta present; apical nematiform setae within fossa at about 0.8× gonocoxite length. Thus the gonocoxal configuration is most like that of M. moorei (Fig. 10A), another early divergent species within subgenus Eucyclothorax (Fig. 1).

Holotype female (ANIC): Walpole N.P. /Zig Zag Rd. WA / 20–27 June 1980 / S. & J. Peck SBP 70B // berlesate / rotted log / litter and / fungi // HOLOTYPE / Mecyclothorax / isolatus / J.K. Liebherr 2018 (black-margined red label).

The adjectival species epithet isolatus signifies both the geographic isolation of this species that is distributed in the south coast region of Western Australia (Fig. 11B), as well as the phylogenetic isolation of this species, as it comprises the adelphotaxon to all other species of subgenus Eucyclothorax (Fig. 1)

This species is known only from the tingle tree (Eucalyptus jacksoni Maiden) forest in the south coast region of Western Australia (Fig. 11B). The single individual was collected in berlesate moist enough to support growth of fungal hyphae (S. B. Peck, pers. comm.). The vestigialized flight wings occur in concert with an extremely abbreviated metathorax and a well-developed plical lock between the elytra and apical abdominal ventrite. This species’ adelphotaxic relationship to the remainder of subgenus Eucyclothorax (Fig. 1) supports its long-term term persistence in this region since the early diversification of all Mecyclothorax.

(n = 5). This species and M. jameswalkeri are the only species of subgenus Eucyclothorax with glabrous hind pronotal angles (Fig. 2E). Of the two, M. darlingtoni is more broad-bodied (Fig. 4A–B), with: 1, a more transverse and basally constricted pronotum, MPW/PL = 1.32–1.35, MPW/BPW = 2.21–2.32; and 2, relatively broader elytra, MEW/EL = 0.70–0.75. Both species are characterized by large eyes, but the eyes of M. darlingtoni do not cover as much of the ocular lobe; EyL/OLL = 0.84–0.88. The parascutellar striole in this species is composed of 4–5 small, deep, isolated pits arcuately joining the basal groove. Standardized body length 4.1–4.8 mm. Setal formula ++/+‒/+2++.

Figure 4. 

Mecyclothorax (Eucyclothorax) spp., dorsal view: A, M. darlingtoni; B, M. jameswalkeri holotype; C, M. lophoides lectotype, Melbourne; D, M. lophoides, ACT: Smoker’s Gap.

Head capsule broad, vertex broadly convex between deep, sinuous frontal grooves, the grooves more shallowly continued onto clypeus; labrum broadly, moderately concave; mandibles moderately elongate, overall length 1.64× distance from anterior condyle to lateroapical labral margin; ocular lobes convexly projected, outer eye surface slightly more convex than posterior portion of lobe meeting gena; mentum tooth with sides acute, apex rounded; ligular apex moderately narrowed, 2 ligular setae separated by 2 setal diameters; paraglossae thin, extended 2× as far past ligular margin as distance from base to ligular margin; mentum breadth/length across lateral lobes = 2.58. Pronotum with articulatory socket of lateral seta touching marginal depression; hind angles obtusely rounded, margin anterad angle slightly concave (Fig. 2E); median base completely margined, the marginal bead uniform and continuous across width (Fig. 4A), base convex anterad basal margin, nearly coplanar with disc though disc is convex and upraised anterad toward middle of pronotum; about 6 minute punctures on base each side of midline, laterobasal depression a shallow longitudinal depression immediately anterad hind angle; median longitudinal impression fine, well indicated, crossed by oblique transverse wrinkles anterad base; anterior transverse impression deep, punctate medially, fine and deep to front angle; anterior callosity slightly convex, defined posteriorly by deep transverse impression; front angle slightly protruded, tightly rounded; prosternal process with broad deep median depression with 4 indistinct pits along its length; prosternum with anteapical groove that is deep and distinctly punctate laterally, smoother and more irregular medially, marginal bead of procoxal cavity bordered anteriorly by distinct, close-set punctures; proepisternum impunctate; proepimeron with broadly raised posterior bead, punctures along suture with proepisternum and anterad posterior bead. Mesepisternum punctate at its deepest portion, about 6 deep punctures in 1–2 dorsoventral rows. Elytra with striae 1–6 composed of isolated punctures in basal half, striae 2–6 reduced to absence in apical 1/3 of length (Fig. 12A), stria 7 absent except near apex from position of subapical sinuation to apical margin; sutural stria deeper in apical half in association with convex sutural interval, the suture smooth from near position of apical dorsal elytral seta to apex (Fig. 12A); stria 8 deep, punctate and bordering lateral marginal depression anteriorly, deeper, smooth and distinct mesad posterior series of lateral elytral setae; lateral elytral setae arrayed in 7 + 6 (anterior series setae and posterior series setae); subapical sinuation evident, abruptly curved anteriorly, but elytral plica covered by margin in dorsal view. Metepisternum moderately elongate, maximum width/lateral length = 0.63; metasternal process with sides acute, apex narrowly rounded, margin very broad medially in apex of process. Abdomen with linear wrinkles on lateral reaches of visible ventrites 1–2, more rounded depressions laterally on ventrites 3–6; suture between ventrites 1 and 2 deeply sinuous laterally; males with 1 seta each side along margin of apical ventrite, females with 2 setae each side and a median patch of 4–5 smaller setae; apical margin of the female apical ventrite with small convex projection medially. Microsculpture absent from frons, the surface glossy, micropunctures visible across the surface; pronotal disc with shallow transverse mesh, sculpticell breadth 3–4× length, pronotal base with evident, regular transverse mesh, sculpticell breadth 2–3× length; elytral disc glossy with fine transverse lines, loosely connected into a mesh and producing an iridescent reflection (Fig. 12A), elytral apex with elongate transverse mesh, surface iridescent; metasternum with distinct transverse mesh, sculpticell breadth 2–3× length; basal abdominal ventrites with swirling transverse mesh and transverse lines, surface iridescent. Coloration of vertex piceous, frons rufous near clypeus, clypeus rufoflavous; antennomeres 1–3 rufoflavous, 4–11 with piceous cast; pronotal disc piceous; margins concolorous; elytral disc dark rufous with iridescent reflection, sutural interval concolorous, interval 9 and marginal depression rufoflavous; elytral apex narrowly brunneous; proepipleuron margin piceous, dark rufous ventrally, proepisternum rufopiceous; elytral epipleura rufoflavous apically, dark rufous ventrally, metepisternum dark rufous; abdominal ventrites rufopiceous with amber posterior margins, apical ventrite with apical half rufobrunneous; femora flavous with brunneous cast; tibiae brunneous with piceous cast.

Male genitalia (n = 4). Male aedeagal median lobe robust, curved, with broad blunt apex, the apical margin curled toward right resulting in a dorsoventral crease (Fig. 7H); internal sac with a robust flagellar sheath, attenuate flagellum, and broad, well-sclerotized dorsal plate (Fig. 7G–H); right paramere elongate, broader basally, conchoid (Fig. 13A), ventral surface with ~9 setae along margin, dorsal margin with 4 setae; left paramere narrow basally, evenly narrowed to apex.

Female reproductive tract (n = 1). Bursa copulatrix elongate, columnar (Fig. 9E); helminthoid sclerite robust with distal projection; spermathecal duct straight, shorter than spermathecal reservoir, evenly sclerotized; basal gonocoxite with 3–4 setae along apical margin, 2 larger laterally and the balance small and positioned medially (Fig. 10E), an apicomedial seta present; apical gonocoxite subtriangular, apex narrowly rounded; lateral ensiform setae small relative to gonocoxite length; apical nematiform setae in apical sensory furrow.

Holotype male (MCZ deposited in ANIC): c30 mi.N.of / Brisbane SQ / Mar. ‘58 / Darlingtons // HOLOTYPE / Mecyclothorax / darlingtoni / J.K. Liebherr 2018 (black-margined red label). Allotypic paratype female (MCZ deposited in ANIC): (same data as holotype, with black-margined red Allotype label).

Paratypes (61 specimens). AUSTRALIA: Queensland: Brisbane, 30 mi. N, iii-1958, Darlingtons (CUIC, 1; MCZ, 19); Mt. Webb, 3 km NE, 15. 03°S 145. 09°E, 03-v-1981, Calder (ANIC, 1); Woondom For. Res., Mothar Mtn. For. Dr., dry rainfor., palm gully, FMHD #2004-217, berl. wet litter along stream, Solidovnikov 1139, 26°15.77’S 152°49.48’E, 380-400 m, 09–xii–2004, Solidovnikov (CUIC, 2; FMNH, 24); FMHD #2004-218, berl. litter under palms, Thayer 1139, 26°15.77’S, 152°49.48’E, 09–xii–2004, Thayer (CUIC, 3; FMNH, 10); Thayer 1139, pyr. fog old logs, 26°15.77’S, 152°49.48’E, 380-400 m, 09–xii–2004, Thayer (FMNH, 1).

The species commemorates Prof. Philip J. Darlington, who collected extensively across Australia during various expeditions undertaken throughout his career. He personally developed the most extensive collection of Australian Carabidae housed in North America, allowing American scientists the ability to work with the fauna. He also curated the Thomas G. Sloane collection after its receipt by C.S.I.R.O., stabilizing the specimens and thereby preserving their information for future researchers. Although he focused on the New Guinea carabid fauna (summarized in Darlington 1971), he rightfully viewed the New Guinean fauna as an extension of the Australian, making biogeographic connections underpinned by taxonomic relationships for much of the Australian Region (Wallace 1876).

M. darlingtoni is broadly distributed along the Queensland coast, with recorded localities spanning the vicinity of Brisbane to Mt. Webb in northern Queensland (Fig. 11C). The extensive numbers of specimens collected in Woondom Forest Reserve (FMNH) were extracted from Berlese samples from mesic litter under palms, or from wet litter along a stream. The single specimen from the northerly and, based on present specimens, disjunct Mt. Webb locality is macropterous, as is one of the two specimens from Dalby, whereas all other specimens from southern Queensland are vestigially winged.

(n = 1). This, the second of two Australian species of subgenus Eucyclothorax– with M. darlingtoni–characterized by the absence of basal pronotal setae, can be diagnosed by aspects of the narrow body, including: 1, a narrow, basally constricted pronotum, MPW/PL = 1.28, MPW/BPW = 1.88; and 2, narrow, subparallel elytra, MEW/EL = 0.64. Like M. darlingtoni, the prosternum of this species has a punctate medial depression anterad the prosternal process, in this instance lined with 7 punctures. The prosternal anteapical groove is continuous and distinctly punctate laterally, more irregular and smoother ventrally. The eyes are large and moderately convex, and they cover much of the ocular lobe; EyL/OLL = 0.95. The parascutellar striole is composed of 7–8 deep, isolated pits. Standardized body length 4.9 mm. Setal formula ++/+‒/+2++.

Head elongate with large eyes (Fig. 4B), vertex broadly convex between deep, sinuously convergent frontal grooves that continue anterolaterally onto clypeus, anterior supraorbital seta in deep depression behind frontolateral callous; labrum broadly, slightly emarginate; mandibles elongate, overall length 1.9× distance from anterior condyle to lateroapical labral margin; mentum tooth with obtuse side, apex shallowly bifid; mentum breadth/length across lateral lobes = 2.57. Pronotum with lateral setal articulatory socket within associated expansion of lateral marginal depression, the lateral marginal depression otherwise broad enough to observe microsculpture at its deepest part, its margin beaded; hind angle obtusely rounded, the lateral margin slightly sinuate before angle; medial base unmargined inside marginal bead that extends only slightly inside hind angle, convex, with ~10 small punctures each side from midline to laterobasal depression; laterobasal depression a narrow oblique groove extended from mesad sinuate lateral margin toward middle of disc; median longitudinal impression very fine and shallow, occluded by broad, shallow transverse wrinkles, deepest between arms of anterior transverse impression; anterior transverse impression smooth, continuous from near midline to rounded, slightly protruded front angles; marginal bead of procoxal cavity lined anteriorly with 3 small isolated punctures: proepisternum impunctate. Mesepisternum punctate at its deepest portion, about 9 deep punctures in 2–3 dorsoventral rows. Elytra with serially punctate striae 1–6, stria 7 absent except near subapical seta; elytral punctures round with distinct center point, separated by their diameter on disc, smaller in stria 6 where separated by 2 punctural diameters; sutural stria distinctly punctate basally, becoming smoother toward apical 1/4 of length, but sutural interval slightly convex (teneral condition?), the transition from punctate to smooth as in Fig. 12A; stria 8 deep, punctate and bordering lateral marginal depression anteriorly, deeper, smooth and distinct mesad posterior series of lateral elytral setae; lateral elytral setae arrayed as 7 + 6 (anterior series setae and posterior series setae); subapical sinuation evident, abruptly curved anteriorly, but elytral plica covered by margin in dorsal view. Metepisternum moderately elongate, maximum width/lateral length = 0.44; metasternal process acute, apex knob-like, margins upraised, twice at broad at apex; metathoracic flight wing macropterous, veins evident in folded condition under teneral elytra, apex reflexed. Abdomen with linear wrinkles laterally on visible ventrites 1–3, shallow rounded depressions laterally on ventrites 3–6; suture between ventrites 1 and 2 slightly sinuate, deep midway along suture; female with 2 marginal setae each side of apical ventrite plus a trapezoidal patch of 4 smaller medial setae. Microsculpture of frons indistinct, surface glossy, vertex with indistinct transverse mesh in parts; pronotal disc and median base with shallow transverse mesh, sculpticell breadth 3–4× length, sculpticells irregularly swirling in laterobasal depressions; elytral disc with transverse lines loosely connected into mesh, the surface subiridescent; elytral apex with transverse lines in a loose mesh, the convex, estriate surface iridescent. Coloration (assessed on single teneral specimen) of vertex rufous with brunneous cast; antennomeres 1–3 rufoflavous, 4–11 with piceous cast; pronotal disc dark rufous, margins paler, flavobrunneous; elytral disc rufobrunneous, sutural interval concolorous, lateral marginal depression rufoflavous, translucent; elytral apex broadly flavobrunneous; proepipleuron rufoflavous, proepisternum rufobrunneous; abdomen rufobrunneous, apical half of apical visible ventrite paler, rufoflavous; femora and tibiae flavous with brunneous cast.

Female reproductive tract (n = 1). The single teneral female specimen of this species was not dissected.

Holotype female (BMNH): Albany / W. Australia / J.J. Walker // G. C. Champion Coll. / B. M. 1927–09 // HOLOTYPE / Mecyclothorax / jameswalkeri / J.K. Liebherr 2018 [black-bordered red label].

This species commemorates James John Walker, Commander and Fleet Engineer, Royal Navy, active member and officer in many scientific societies–including President of the Linnean Society of New South Wales (Walker 1921)–and in retirement, an editor of the Entomologistʼs Monthly Magazine (Poulton 1939). Walkerʼs collections from Australia and New Zealand were passed to George C. Champion, his brother-in-law, and then bequeathed by Champion to The Natural History Museum, London. Walkerʼs collections were as far flung as the British Empire, with his naval duties taking him to places where he could collect. Among many other species, carabid beetles named after him include: Protopaussus walkeri Waterhouse (1897) from China; Calosoma walkeri Waterhouse (1898), junior synonym of C. oceanicum Perroud, from Australia; Rhaebolestes walkeri Sloane (1903) from New South Wales; Duvaliomimus walkeri (Broun 1903), Megadromus walkeri (Broun, 1903), and Mecodema walkeri Broun (1903), the last a junior synonym of Mecodema howitti (Castelnau 1867), all from South Island, New Zealand; and Trirammatus walkeri (Andrewes 1931) from Juan Fernandez Island. Walkerʼs collecting acumen can be attested to by his discovery of this broad assortment of carabid diversity.

The lone specimen of this species is from Albany, W.A. (Fig. 11C). We know nothing specific about the habitat in which this beetle was discovered. The specimen is macropterous with the wings bearing well-developed venation and a reflexed apex, the wing structures being visible through the very translucent elytra of the teneral specimen. Thus occupation of a riparian habitat requiring recolonization of habitat patches may be predicted (Darlington 1936, 1943).

(n = 5). This species is characterized by a narrow, moderately cordate pronotum, the lateral margins slightly sinuate anterad obtuse, moderately projected hind angles (Fig. 2F); MEW/BPW = 1.57–1.68, MPW/PL = 1.18–1.22. The pronotal lateral marginal depression is very narrow, with only the narrowest indication of microsculpture between the convex disc and the marginal bead. The hind angle is obtuse, rounded at its apex, with the basal pronotal seta set slightly anterad the angle. The marginal depression continues for only a short distance mesad the hind angle. The median base is covered with ~10 erratically distributed small punctures each side of midline. The laterobasal depression is a longitudinal depression lined with 4–5 larger punctures, with the flat area between depression and the marginal bead also bearing several larger punctures. The prosternum is medially depressed from the prosternal process between the procoxae 1/2 the distance toward the anterior margin, the depression lined with 6–7 pits. The anteapical groove is shallowly punctate laterally, continuous and irregularly indented medially, and the marginal bead of the procoxal cavity is bordered anteriorly by about 3 very shallow punctures. The mesepisternum is punctate at its deepest portion, about 9 deep punctures in 2–3 dorsoventral rows. The elytra are narrowly subparallel (Fig. 4C–D), with the humeral angles distinctly obtuse; MEW/EL = 0.64–0.69. The parascutellar striole is composed of 4–6 small, deep, isolated punctures. Elytral striae 1–6 are present on the disc, though striae 2–6 are absent basally and from the apical quarter to half, the outer striae progressively shorter. The strial punctures are isolated, progressively so apically, and the sutural stria is smooth or only slightly punctate in the apical half (Fig. 12B). Body coloration varies from dark brunneous (Fig. 4C) to piceous (Fig. 4D), with the legs correspondingly brunneous to piceous; i.e. there is less contrast between leg and body color than in M. peryphoides (Fig. 5B) or M. cordicollis (Fig. 5C). The elytral apex may be slightly paler than the disc in the brunneous specimens, however any difference is gradual, not a distinct transition as in M. cordicollis (Fig. 5C). Elytral margins are concolorous with the disc in the darker specimens. Cuticular microsculpture is relatively less developed in this species than in M. darlingtoni, M. jameswalkeri, M. peryphoides, or M. cordicollis, with: 1, frons glossy, indistinct transverse lines visible over portions of the surface; 2, pronotal disc glossy with shallow elongate transverse microsculpture visible outside the area of reflection; 3, flat elytral intervals covered with dense transverse lines producing an iridescent reflection. Standardized body length 3.8–4.9 mm. Setal formula ++/++/+2++.

Male genitalia (n = 12). Aedeagal median lobe dorsoventrally broad, the apex broadly rounded and slightly projected beyond the apical margin of ostium (Fig. 14A, C); flagellum elongate and hooklike, the flagellar sheath of similar length, its surface scabrous (Fig. 14A–B), dorsal plate ovoid, lightly sclerotized (Fig. 14B); right paramere expanded basally, narrowed beyond midlength, the ventral surface with ~9 setae along margin, dorsal margin with 4 setae in apical half (Fig. 13B); left paramere narrow basally, evenly narrowed to apex.

Female reproductive tract (n = 2). Bursa copulatrix squat, as broad as long (Fig. 9F); helminthoid sclerite broad basally, with distinct mediodistal projection; spermathecal duct straight, narrow, evenly sclerotized, as long as spermathecal reservoir; basal gonocoxite with 4–5 apical setae plus a large seta at the apicomedial angle (Fig. 10F); apical gonocoxite subtriangular, broadly rounded apically; lateral ensiform setae small, narrow; apical nematiform setae in apical sensory furrow.

Lectotype male (MNHN) hereby designated: pointed specimen // Musaeo Chaudoir [red typeface] // Lectotype [red label] // Museum Paris / ex. Coll. Oberthur // Mecyclothorax (Ch.) det. P. M. Johns. There are also a male paralectotype and a female paralectotype (MNHN).

M. lophoides is allopatrically distributed to the south of its adelphotaxon M. darlingtoni, with localities ranging from northeastern New South Wales southward through eastern N.S.W. to Melbourne (Fig. 11C). Non-type material and repositories include: A.C.T.: Paddy’s R. 1 mi. S Cotter Dam (ANIC, 1); Smoker’s Gap 43 km SW Canberra (CAS, 10; CUIC, 2): NSW: Blackheath (MVM, 1); Braidwood (CUIC, 2; MCZ, 37); Mt. Kosciuszko (MCZ, 2); New England N. P., Thungutti Camp (ANIC, 3): VIC: Dandenong Ck. (MVM, 1); Gellibrand R., Otway Ranges (MCZ, 1); Oakleigh (MVM, 1); Portland to Pt. Fairy (CUIC, 1; MCZ, 21); Wilson’s Promontory (ANIC, 1); Winchester (MCZ, 1).

Specimens of this species collected by John Nunn on King Island (Moore 1984: 164) were preceded temporally by beetles laid down from 143,000–75,000 years ago in subfossiliferous deposits at Yarra Creek, King Island, during the Pleistocene last interglacial (Porch et al. 2009). Long-term residence on King Island suggests the species can persist in communities ranging from the present more mesic, more seasonal forest types to the wetter, more aseasonal forests present on King Island during the Pleistocene. All specimens are vestigially winged.

(n = 5). Among species of the M. lophoides complex this species stands out based on its rufous coloration (Fig. 5A), and distinctly cordate pronotum with projected, nearly right hind angles (Fig. 2G). In the most melanized specimens, the forebody–head and pronotum–may be rufopiceous, but the elytra retain rufous coloration, and the legs are pale, flavous, with a slightly smoky piceous cast. The eyes are moderately convex, MHW/mFW = 1.51–1.57, less convex than those of the other M. lophoides complex species (Figs 4, 5B–C). The pronotum is quite constricted basally, MEW/BPW = 1.76–1.82, and moderately transverse, MEW/PL = 1.22–1.28. The pronotal lateral marginal depression is moderately broad, with the marginal bead only slightly upraised. The median base is minutely punctate, with 5–8 isolated punctures each side from midline to the ill-defined laterobasal depressions, those defined mostly by a longitudinal line of larger punctures, with several larger punctures also present between that line and the marginal bead. The hind angle is right to slightly obtuse, with the posterior margin transverse and anterad the convex median basal margin, which is smooth not beaded. The prosternum is flat to depressed medially, the medial area bearing a longitudinal series of 7 distinct punctures. The anteapical groove is deep and distinctly punctate laterally, continuous and more shallowly punctate medially, and the marginal bead of the procoxal cavity is bordered anteriorly by 5–6 strigose punctulae. The mesepisternum is covered with punctures, about 13 deep punctures arrayed in 3–4 irregular rows. The elytra are relatively broad, MEW/EL = 0.67–0.73, and flat medially on disc. Elytral striae 1–4 bear large punctures, those serial punctures close set enough on the disc to depress the intervening cuticle. Strial punctation is more strongly developed in this species than in other species of the M. lophoides complex (Fig. 12), with the sutural stria distinctly punctate mesad the posterior dorsal elytral seta. In this species cuticular microsculpture is less well developed than in the other M. lophoides complex species, with: 1, frons smooth, glossy, with micropunctures sporadically visible across surface; 2, pronotal disc glossy with sporadic micropunctures visible, very transverse lines sporadically visible laterally and near concavities of median longitudinal impression and laterobasal depression; 3, elytral discal intervals largely glossy, with shallow transverse lines sporadically visible over surface (Fig. 12C), transverse microsculpture more developed on elytral apex where it forms an elongate transverse mesh, sculpticells 3–4× broad as long. Body coloration varies from a bright rufous, mostly in desert inhabiting beetles from the northern part of the range including the type locality of Leighʼs Creek (Fig. 11D), to darker with rufopiceous head and pronotum and rufous elytra at localities on the southern edge of the range; Mt. Remarkable and Telowie Gorge. Even in the darker specimens, the base of the pronotum is an amber rufous, the cuticle appearing translucent. Standardized body length 4.2–5.5 mm. Setal formula ++/++/+2++.

Male genitalia (n = 10). Aedeagal median lobe broadest near basal 1/3 of length, ventral margin distinctly and evenly curved (Fig. 14D); internal sac bearing a long sinuous flagellum, a large bean-shaped dorsal plate, and a well-sclerotized, smooth flagellar sheath, the sheath length less than 1/4 flagellar length (Fig. 14D, as in Fig. 14F); right paramere expanded dorsally in basal half, the dorsal margin convex, 7–11 setae along ventral margin and ~1 on dorsal margin (Fig. 13C); left paramere broadly quadrate basally, extended as a narrowly attenuated whip to apex. Comparing the dissected aedeagi of 10 M. eyrensis males (e.g. Fig. 14D) and 16 M. peryphoides males (e.g. Fig. 14E) resulted in no discernible differences in overall shape of the median lobe, in curvature or expanse of the apex, in the basal bulb or development of the sagittal crest, nor in structures of the internal sac such as the flagellum, flagellar sheath, or dorsal plate (Fig. 14D–E). The only male genitalic differences noted between males of these two species involved the presence of fewer setae along the ventral margin of the right parameres of M. eyrensis versus M. peryphoides (e.g. Fig. 13C–D). To determine whether these differences could diagnose the species, the numbers of setae along the ventral margin of the right paramere in 10 individuals of M. eyrensis (setal numbers: 7, 7, 8, 8, 8, 8, 9, 11, 11, 11) were compared to those in 15 individuals of M. peryphoides (setal numbers: 9, 10, 10, 11, 11, 11, 12, 12, 13, 13, 13, 14, 15, 15, 17) using the Wilcoxon rank-sum test (Snedecor and Cochran 1980). The distribution of values resulted in a minimal T₁ = 69, below the threshold value of 84 for a p = 0.01 level of significance for difference between the two distributions (Supplementary material 2). Even though the values among individuals overlap slightly in the 9–11 setal counts, there is a statistically significant difference between the setal configurations of the two species. This significant difference augments the diagnostic differences found in the external characters.

Female reproductive tract (n = 3). Bursa copulatrix short, slightly longer than broad (Fig. 9G); helminthoid sclerite broad basally, with narrow, elongate mediodistal projection; spermathecal duct elongate, sclerotized enough to hold coiled configuration, length ~2× length of spermathecal reservoir; basal gonocoxite with 2–6 apical setae, 2–4 of those larger and the balance smaller, plus a similarly sized seta at the apicomedial angle (Fig. 10G); medial surface of basal gonocoxite with several smaller setae along length; apical gonocoxite subtriangular, narrowly rounded apically; lateral ensiform setae small, narrow; apical nematiform setae in apical sensory furrow.

Figure 5. 

Mecyclothorax spp., dorsal view: A, M. (Eucyclothorax) eyrensis, paralectotype; B, M. (Eucyclothorax) peryphoides; C, M. (Eucyclothorax) cordicollis; D, M. (Qecyclothorax) lewisensis estriatus holotype.

Lectotype male (BMNH) hereby designated: platen mounted with “2710 T / L. C. (red ink) // Type (round, red-margined label) // Blackburn coll. 1910-236 // Cyclothorax eyrensis Blackb. // Lectotype Cyclothorax eyrensis Blackburn / J.K. Liebherr 2006 [black-margined red label]. The abbreviation “L. C.” stands for Leigh Creek based on the labels below. Paralectotype female (SAMA): platen mounted with L. C. 2710 in red ink // Leigh Ck. C.A. / Blackb’s Coll. // Cyclothorax eyrensis Bl. / Co-type // Mecyclothorax eyrensis Blkn / S. Australia / Cotype [red ink] // Paralectotype ♀ / Cyclothorax / eyrensis Blackburn / det. J.K. Liebherr 2004 [black-margined red label]. The type locality is Leigh Creek, South Australia.

This species is distributed across the interior of southeastern Australia (Fig. 11D), in South Australia from Telowie Gorge to Mt. Remarkable (FMNH), eastward to western N.S.W., and northward into southern Northern Territory at Palm Valley and Stokes Creek in the Gill Range (FMNH). Localities and repositories for non-type material include: NSW: Silverton (ANIC, 1). NT: Gill Range, Stokes Ck. (CUIC, 1; FMNH, 2); Palm Valley (FMNH, 1). SA: Flinders Ranges, Bunyeroo Gorge (CUIC, 1; FMNH, 14), Telowie Gorge (CUIC, 2; FMNH, 5); Mann Range, 0.5 km N Angatja Homestead (CUIC, 1; FMNH, 1); Musgrave Range, 13 km N Ernabella (CUIC, 1; FMNH, 10), 15 km W Officer Ck. (FMNH, 10), 17 km W Jacky Pass (CUIC, 1); Mt. Remarkable (CUIC, 6; FMNH, 94); Wilmington (MVM, 3); Yunta (FMNH, 3). The extensive collections made by L. Watrous (FMNH) were obtained from sieved litter or reed drift from along dry or running streams. Nonetheless, and going against Darlingtonʼs generalization for riparian species, all observed individuals are vestigially winged.

(n = 5). Among the species of the M. lophoides complex characterized by darker bodies and contrastingly pale legs (Figs 4B, 5B–C), this species has an ellipsoid elytra with large strial punctures, a pronotum with distinct obtuse hind angles, and concolorous elytral disc and apex, i.e. without an apical flavous marginal band (Fig. 5B). The eyes are broadly convex, MHW/mFW = 1.60–166. The pronotal margins are sinuate anterad the well-defined hind angles, but the posterior margin behind the angle is at most slightly sinuate, and usually convex, smoothly meeting the curved median basal margin (Fig. 2H). Body coloration is dark, with the head, pronotum and elytra piceous. The pronotum is transverse and constricted basally–MEW/EL = 1.20–1.28, MPW/BPW = 1.63–1.71–though the basal constriction less than that observed in beetles of M. eyrensis; MEW/BPW = 1.76–1.82. The pronotal median base is covered with distinct punctures, about 20 punctulae each side from midline to the very shallow, poorly defined laterobasal depression (Fig. 2H). The pronotal lateral marginal depression is narrow, but broad enough for sculpticells to line the deepest part, and the margin is beaded. Like M. eyrensis: 1, the prosternum is flat to depressed medially, the medial area bearing a longitudinal series of 4–7 distinct punctures; 2, the anteapical groove is deep and distinctly punctate laterally, continuous and more shallowly punctate medially; 3, the marginal bead of the procoxal cavity is bordered anteriorly by 5–6 strigose punctulae; and 4, the concave mesepisternal surface is lined with 13 deep punctures arrayed in 3–4 irregular rows. The elytral striae are punctate on the disc, though the punctures are far enough apart that the cuticle is coplanar with the intervals between the punctures. Like M. eyrensis, the punctures of striae 1–6 are progressively more isolated laterally on the elytra, but the sutural stria is more shallowly and irregularly punctured mesad the posterior dorsal elytral seta in this species (Fig. 12C–D). Cuticular microsculpture is well developed in this species, with: 1, frons covered with indistinct transverse lines, vertex with more well-developed sculpticells, transforming from transverse just behind posterior supraorbital setae to nearly isodiametric near pronotum; 2, pronotal disc with indistinct transverse lines visible outside areas of direct reflection, irregular surface of pronotal base covered with irregular transverse mesh; 3, elytral disc with transverse lines visible outside areas of direct reflection, apex covered with well-developed elongate transverse mesh. Standardized body length 4.3–5.0 mm. Setal formula ++/++/+2++.

Male genitalia (n = 16). As stated under M. eyrensis, the aedeagal median lobe and flagellar complex of that species and M. peryphoides show no differences (Fig. 14D–E). However the right paramere of this species bears significantly more setae (9–17) along its ventral margin.

Female reproductive tract (n = 3). Bursa copulatrix short, slightly longer than broad (Fig. 9H); helminthoid sclerite broad basally, with narrow mediodistal projection; spermathecal duct elongate, sclerotized enough to hold coiled configuration, length ~2× length of spermathecal reservoir; basal gonocoxite with 3–6 apical setae, 2–3 large and the balance smaller, plus a similarly sized seta at the apicomedial angle (Fig. 10H); medial surface of basal gonocoxite with larger setae apically; apical gonocoxite subtriangular, narrowly rounded apically; lateral ensiform setae small, narrow; apical nematiform setae in apical sensory furrow.

Holotype male (BMNH): platen mounted with “1614” in red ink and “T” in black ink on the obverse, with basal marginal black and red lines, the red crossed by an arrowhead // Type (round red-margined label) // Blackburn coll. 1910-236. // Cyclothorax periphoides [sic.], Blackb. Blackburn states the type locality as “Woodville, near Adelaide; a single specimen (Blackburn 1889: 1393).”

This species is distributed (Fig. 11D) to the south of its sister species, M. eyrensis, in South Australia. The type locality of Woodville is near recent collections made at Belair Recreation Park, 10 miles S of Adelaide (FMNH), Blackwood, Sturt Gorge Reserve (ZMUC), and Hale Conservation Park SE Williamstown (FMNH). To the east the species is recorded from the following localities: A.C.T.: Black Mountain (CUIC, 8; EMEC, 1); NSW: Belmore (AMS, 1); Bogan R. (AMS, 1); “Calosoma” via Gundaroo (ANIC, 4); Federal Hwy. (ANIC, 1); Jerrawa (AMCS, 2); Mt. Wilson (CUIC, 1; FMNH, 4); Narrabeen (AMS, 1); Queanbeyan (ANIC, 1); Tuglo Wldlf. Ref. 48 km N Singleton (FMNH, 1); Weddin Mtns. N. P. (FMNH, 1); VIC: Lake Eildon N. P., Sebastopol Ck. (ZMUC, 1). Collections have been made via litter sifting along streams or in Eucalyptus (Myrtaceae) or mixed deciduous forest, or in rainforest on Mt. Wilson. As for M. eyrensis, even though this species occupies predominantly riparian situations, it appears to be vestigially winged.

(n = 5). This species (Fig. 5C) and M. lophoides (Fig. 4C–D) include the smallest-bodied, most gracile beetles in the M. lophoides species complex, with standardized body length for this species = 4.0–4.7 mm. Individuals exhibit a flavous marginal band on the elytra, brunneous to rufobrunneous body, and pale, flavous legs. The pronotum is very similar to that of M. lophoides (Fig. 2F, I); moderately transverse, MPW/PL = 1.17–1.24, and basally constricted, MPW/BPW = 1.62–1.68, these values broadly overlapping those calculated from specimens of M. lophoides. The eyes are also similar in configuration in the two species: here MHW/mFW = 1.63–1.71, with the eyes covering most of the ocular lobe, EyL/OLL = 0.86–0.91. The pronotal lateral marginal depression is slightly broader in this species than in M. lophoides, with microsculpture evident in its deepest part. The pronotal hind angles are obtusely rounded, with the margin behind the angle smoothly curved onto median base. The marginal bead terminates at the laterobasal depression just mesad the hind angle, and the bead is only slightly broader at the setal insertion. The pronotal median base is covered with 10–13 large isolated punctures present each side of midline. The laterobasal depression is defined by a medially arcuate line of punctures commencing at the termination of the marginal bead mesad the basal seta. The prosternum is medially depressed from the prosternal process anterad slightly less than 1/2 the distance to the anterior margin, the depression broader anteriorly, and lined with ~8 pits. The anteapical groove is punctate laterally, continuous and indistinctly punctate medially, and the procoxal cavity marginal bead is bordered by 4–5 small punctures along its anterior margin. The mesepisternum is variously punctate, the deepest portion covered with about 9 shallow to deep punctures arranged in 2–3 dorsoventral rows. The elytra are broad basally, subparallel, and broad relative to their length (Fig. 5C), MEW/EL = 0.69–0.71, versus values of 0.64–0.69 for individuals of M. lophoides (Fig. 4C–D). The elytral disc is flat, with the sutural intervals apically raised into a callus. Elytral striae 1–6 are distinctly punctate on the disc, with the punctures nearly contiguous on the inner striae, however the intervening intervals are nearly flat. As in the other M. lophoides complex species, strial punctures are smaller and therefore further apart along the lateral striae, though in this species stria 6 is indicated until posterad midlength. As in M. peryphoides, the sutural stria is indistinctly punctate mesad the posterior dorsal elytral seta (Fig. 12D–E), but the punctures in striae 2–4 are more well developed, agreeing with M. eyrensis, but not M. lophoides (Fig. 12 B–C). Coincident with the presence of a flavous marginal band on the darker elytra, other body coloration tends toward the polychromatic: 1, elytral epipleura dorsally flavous versus ventrally rufobrunneous adjacent to the metepisternum; 2, coxae dark brunneous, trochanters rufous, femora flavous, and tibiae and tarsomeres brunneous with smoky cast. Cuticular microsculpture includes: 1, vertex glossy with an indistinct transverse mesh in parts; 2, pronotal disc glossy with an elongate transverse mesh restricted to transverse wrinkles, and a shallow elongate transverse mesh on the median base, sculpticells more irregular in the laterobasal depression; 3, elytral disc with a shallow, elongate transverse mesh, sculpticell breadth 3–4× length, the convex apex covered with traceable transverse lines, the entire elytral surface subiridescent. Setal formula ++/++/+2++.

Male genitalia (n = 13). Aedeagal median lobe of moderate dorsoventral breadth (Fig. 14G), evenly curved as in M. lophoides (Fig. 14A); flagellar complex including a very short, spine-like flagellum, a bifurcated flagellar sheath, and a lightly sclerotized, ovoid dorsal plate (Fig. 14H); right paramere broader basally, conchoid, with ~12 setae along ventral margin and 2–3 small setae on dorsal margin (Fig. 13E); left paramere broadest basally, but evenly narrowed to whiplike apex, the apex flexible and often twisted relative to base when aedeagus mounted on slide.

Female reproductive tract (n = 1). Bursa copulatrix squat, as broad as long (Fig. 9I); helminthoid sclerite broad basally, with narrow elongate mediodistal projection; spermathecal duct straight and narrow, evenly sclerotized, of similar length to spermathecal reservoir; basal gonocoxite with 2–3 apical setae, 2 larger setae laterally, and a moderately sized seta at the apicomedial angle (Fig. 10F); apical gonocoxite subtriangular, narrowly rounded apically; 1–2 small lateral ensiform setae present; apical nematiform setae in apical sensory furrow.

Lectotype male (ANIC) hereby designated: 2nd specimen from left on 6-specimen platen, specimen annotated “♂ -w”, male genitalia dissected and placed in polyethylene genitalia vial below labels // Cyclothorax Type / cordicollis Sl. MSS. / Colombo Plains 11/6/95 // C. cordicollis Sl. / Holotype / PJD not HT JKL 18 [pink label with Holotype crossed out] // ANIC Database / 25 014958 // ANIC Specimen [green label] / ANIC Image [orange label] // Lectotype / Cyclothorax / cordicollis / J.K. Liebherr 2017 [black-margined red label] / Mecyclothorax / cordicollis / (Sloane) ♂LT / det. J.K. Liebherr 2018. Although P.J. Darlington labelled these specimens as “Holotype”, a lectotype must be designated given Sloaneʼs description stating the multiple localities given as “Queensland–Brisbane (sent by Mr. Lea); N.S. Wales–Clarence River and Windsor (Lea), Grenfell, Junee, Urana, Mulwala (Sloane); Victoria–Ferntree Gully and Lilydale (Sloane) (Sloane 1900: 564).” Columbo Creek [Colombo Plains sic] is near and northwest of Lake Urana and the town of Urana, and so these specimens would correspond to the “Urana” locality of Sloaneʼs list. Moreover, Sloane wrote regarding the habits of these beetles; “Found in damp situations near water, usually rare, but on July 11th, 1895, I found it very plentifully under sticks along the edge of a swamp about 20 miles north from the town of Urana (Sloane 1900: 564).” As the label clearly reads “11/6/95” and Sloane wrote of “July 11th, 1895”, some ambiguity concerning the date of this collection remains [though the label data would hold precedence]. However no such ambiguity can be countenanced regarding this collecting locality.

Sloane (1900) described this species from specimens representing much of the distributional range as now known augmented by more than another century of collections (Fig. 11E), with the single notable exception of two specimens collected by the Darlingtons, ii–1958, at Ravenshoe, W of the Atherton Tableland (MCZ, 2). Localities and repositories of other non-type specimens include: NSW: Bodalla (CUIC, 1; MCZ, 10); Gosford (MVM, 2); Narrandera (MCZ, 1); Sydney vic. (CUIC, 1; MCZ, 15); QLD: Blunder (ANIC, 1) ; Brisbane vic. (CUIC, 2; MCZ, 9); Dalby (ANIC, 2); VIC: Dandenong Vy. (ANIC, 2); Lilydale (ANIC, 1); Melbourne (ANIC, 1); Powlett R. (ANIC, 1).

Sloaneʼs perspicacity with regard to species boundaries is very evident in his sorting out this taxon from the very similar appearing beetles of M. lophoides and M. peryphoides. What we know about the habits of this species can be taken from Sloaneʼs description, quoted above. The species is uniformly represented by vestigially winged individuals with the exceptions of macropterous beetles from two Queensland localities: 1, one of two beetles from Ravenshoe; and 2, two of two specimens from Dalby.

These robust-bodied species (e.g. Fig. 5D) are geographically restricted to Queensland, Australia, and have been recently revised by Baehr (2003). The pronotum of species in this subgenus is broad, with obtuse or obtusely rounded hind angles. Each elytron bears a single dorsal elytral seta just before midlength (Fig. 5D). The prosternum is medially depressed both between and anterad the procoxal cavities, a condition shared with most member species of Eucyclothorax, although punctures are not present in the depression. The elytral striae are reduced; striae 1–3 to 1–4 shallow to evanescent, striae 4– or 5–7 obsolete. Striae 1, 7, or both may be present apically on the elytra, but striae 4–6 are consistently absent there. The elytra are broadly convex, with interval 8 not, or only slightly upraised (M. lewisensis) relative to the general curvature of the elytral surface. The suture between abdominal ventrites 1 and 2 is nearly straight, with the second ventrite hardly depressed relative to the first. Body size is small for the genus; standardized body length 2.6–3.7 mm.

The male aedeagal median lobe internal sac bears a flagellum (Liebherr 2018, fig. 4E), and the ventral paramere is elongate-conchoid in shape, broadly to narrowly subtriangular with ventral setae present (Liebherr 2018, fig. 5B; Baehr 2003, fig. 1). The female bursa copulatrix is relatively short (Liebherr 2018, fig. 6D). The spermathecal duct joins the bursa copulatrix-common oviduct juncture.

Baehr (2003) recognized four species, three of them represented by two subspecies each, for a total of seven specific and subspecific forms. The subspecies described below is added to the inventory of M. lewisensis, resulting in three subspecies; M. lewisensis, M. lewisensis uncinatus, plus the newly described subspecies.

(n = 1). This taxon is distinguished from all others of subgenus Qecyclothorax by the reduced elytral striation, with only the sutural stria evident, and the positions of all outer striae only traceable by the longitudinal tracks of trachea (Shelford 1915). As for all Qecyclothorax, the elytra exhibit only a single dorsal elytral seta positioned near midlength (Fig. 5D). Consistent with membership in M. lewisensis, this beetle exhibits a quadrisetose clypeus and a pronotum with explanate lateral margins that are indistinctly sinuate basally. The parascutellar seta are very short and fine, and they are set in shallow depressions, however careful examination allows their discernment along with the fine articulatory sockets from which they extend. Baehr used the comparative breadth of the pronotal base relative to its apex as one criterion to differentiate the two subspecies M. lewisensis and M. l. uncinatus: the former with a ratio APW/BPW < 0.83 (note inverted ratio herein versus Baehr 2003), the latter with APW/BPW > 0.85. In the single specimen of M. l. estriatus, APW/BPW = 0.85. Standardized body length for the type specimen below, 3.3 mm. Setal formula ++/++/+1++.

Head broad, frontal groove deep, arcuately convergent toward clypeus, continued onto clypeus, terminated posteriorly midway between 2 supraorbital setae; eyes moderately convex, MHW/mFW = 1.52, covering much of ocular lobe, EyL/OLL = 0.83; antennae elongate, long enough so that apex would extend to basal 1/4 of elytra, antennomere 9 length/maximal breadth = 1.89; mentum tooth with sides acute, apex broadly rounded; ligular apex narrowed, slightly concave between ligular setae, setae separated by 2 setal diameters; paraglossae extended as far beyond ligular apical margin as distance from base to ligular margin. Pronotum transverse, MPW/PL = 1.41, moderately constricted basally, MPW/BPW = 1.35; lateral pronotal seta placed 1 diameter mesad lateral margin depression, depression very narrow at front, gradually widened to explanate at hind angle; basal margin nearly straight, slightly convex between laterobasal depressions, margin flat and effaced behind laterobasal depressions, a convex roll medially; median base depressed relative to disc, smooth with ~3 small punctures each side mesad laterobasal depression; laterobasal depression a linear to slightly outwardly arcuate line of 3–4 broad punctures, the area laterad line of punctures broadly convex to explanate lateral margin; median longitudinal impression fine, deep, adjacent depression covered with transverse wrinkles on disc; anterior transverse impression broad, evenly depressed fore and aft, the anterior callosity broadly, slightly convex to front margin; front angles slightly protruded, subangulate with marginal bead mesad angle that is continuous with transverse impression; prosternum depressed medially anterad procoxal cavities, the depressionʼs surface irregular with 3 shallow irregularities disturbing the surface; anteapical groove very shallow laterally discontinuous, not present medially; lateral reaches of prosternum irregular, procoxal cavity with very fine marginal bead. Mesepisternum covered with 7 large, isolated pits on a smooth surface, the pits arranged in 2 dorsoventral rows; metepisternum nearly quadrate, lateral margin length 1.2× maximal width. Elytral broadly hemiovoid (Fig. 5D), convex laterally with sides nearly vertical, disc flattened; basal groove slightly curved laterad position of parascutellar striole, punctate near basal positions of striae 3 and 4 (indicated by tracheae; Shelford 1915), and straight laterally to obtusely angulate humeri; elytral striae obsolete, striae 1 and 2 traceable only by very small serial punctures on disc, stria 3 less easily traced as punctures are irregular and easily confused with micropunctures scattered over cuticle; dorsal elytral setae short, in depressions that span only 1/4-1/2 of third interval; only stria 7 evident on elytral apex as a broad shallow depression connecting the subapical and apical elytral setae; stria 8 deep, present from posterad anterior series of lateral elytral setae, slightly irregular along length behind anterior series of lateral elytral setae; lateral elytral setae arranged as 7 + 6–7 setae; subapical sinuation very broad and shallow, the elytral plica visible in dorsal view. Head capsule with reduced microsculpture, frons glossy and vertex covered with fine transverse lines; pronotal disc glossy, a transverse mesh present over parts, sculpticell breadth 2–3× length; pronotal base glossy, indistinct transverse mesh in laterobasal depression; elytral disc glossy, indistinct transverse lines outside area of reflection, apex glossy with indistinct transverse mesh in irregularly depressed areas. Coloration of head rufous on frons, darker on vertex; antennomeres 1 rufoflavous, 2–11 brunneous; Pronotal disc dark rufous, lateral margins broadly paler, rufoflavous, base a translucent amber; elytral disc rufobrunneous overall, but with a darker transverse field posterad dorsal elytral seta that continues along suture to apex leaving 2 paler lateroapical fields; sutural interval concolorous with adjacent intervals; proepipleuron rufoflavous with darker explanate margin; proepisternum dark rufous with subiridescent reflection; elytral epipleuron broadly rufoflavous, metepisternum rufobrunneous; abdomen rufobrunneous, apical 2/3 of apical ventrite flavous; femora flavous, tibiae flavous with rufous cast.

Female reproductive tract (n = 1). The unique female holotype was not dissected. Nonetheless, the gonocoxae extend from the abdominal apex, allowing the following characters to be assessed: basal gonocoxite with medioapical margin glabrous; apical gonocoxite broad basally with 2 lateral ensiform setae; apical nematiform setae in subbasal sensory furrow. These characters conform to states previously scored for M. lewisensis (Liebherr 2018) though they are not definitive.

Holotype female (QMB): QLD: 20°21'S, 148°43'E / Brandy Creek, 150 m / 20 Nov 1992 / Monteith, Thompson / & Janetzki, Pyrethrum // HOLOTYPE / Mecyclothorax / lewisensis / estriatus / J.K. Liebherr 2018 (black margined red label).

The lone specimen of this subspecific taxon is from near the Queensland coast south of Cannonvale (Fig. 11E), approximately 600 km south of the localities near Mossman from where the other subspecies of M. lewisensis have been described (Baehr 2003, fig. 6).

This subgenus comprises over 350 species (Liebherr 2018), and so morphological variation among the species is the most extreme observed among the various Mecyclothorax subgenera. However, throughout taxa of this subgenus, the labrum is emarginate apically, either distinctly and angularly as in M. goweri Moore of Lord Howe Island (Liebherr 2018, fig. 1A), or less so as in M. aeneipennis Liebherr of Haleakalā, Maui, Hawaiʻi (Liebherr 2015, fig. 7). The ligular margin is generally truncate with the ligular setae well separated (Liebherr 2018, fig. 1G), though as exceptions, the ligula is apically rounded in the Papuan taxa M. brispex and M. andersoni (Liebherr 2017, 2018, fig. 7). The prosternum exhibits a smooth to distinctly punctate anteapical groove, though never any other punctures, such as in the median depression observed among species of subgenus Eucyclothorax. The parascutellar striole is present, and may be smooth or punctate, with up to 8 punctures along its length (Fig. 6). Among the mainland Australian species, the pronotal median base is depressed relative to the disc, and punctate; a distinction noted by Blackburn (1889).

The male aedeagus has an internal sac with an apical flagellar plate surrounding the gonopore (Fig. 15E). The female reproductive tract most often has the spermathecal duct entering the bursa copulatrix mediodorsally (Fig. 17C–D), however the sister-species pair M. lateralis + M. minutus (Fig. 1A) revert to the plesiomorphic configuration wherein the spermathecal duct basally joins the juncture of the common oviduct and the bursa copulatrix (Fig. 17A–B).

This subgenus is represented by four species in mainland Australia. Numbers of taxa in the substantial radiations from Hawaii, the Society Islands, New Guinea, New Zealand, the Sundas, and Lord Howe, Norfolk, and St. Paul and Amsterdam Islands are summarized in Liebherr (2018: 15).

Figure 6. 

Mecyclothorax (s. s.) spp., dorsal view: A, M. lateralis; B, M. minutus; C, M. ambiguus; D, M. punctipennis.

(n = 5). This large-bodied species–standardized body length 5.2–6.4 mm–is further distinguished by the rufous to brunneous body with contrasting, flavous elytral margins (Fig. 6A). The labrum is broadly and distinctly emarginate apically. The ligula is truncate apically, with the 2 apical setae separated by 4 setal diameters, the ligular surface longitudinally depressed between the setal articulations. The paraglossae are elongate, apically extended beyond the ligular apical margin twice the distance from ligular margin to their base. The pronotum is robust, transverse, with a convex disc, and depressed and circularly punctate median base (Fig. 2J); MPW/BPW = 1.45–1.56, MPW/PL = 1.35–1.40. The prosternal process is broad, only slightly depressed between the coxae, and convex anterad the coxae. The prosternal anteapical groove is deep and narrow, with only slight irregularities along its length at its deepest part. As in its sister species M. minutus, the parascutellar seta is absent. The metepisternum is abbreviated, with lateral length about 2× maximum width, and the metathoracic wings are vestigial in examined material. The suture between visible ventrites 1 and 2 is sinuous, with the area behind markedly depressed. Microsculpture of the head is reduced, with frons glossy, vertex with indistinct transverse sculpticells; pronotal disc and median base glossy, indistinct transverse lines visible in places; elytral surface glossy with well-defined isodiametric sculpticells visible over entire surface, the apex with sculpticells more upraised; thoracic ventrites glossy, abdominal ventrites glossy medially but with isodiametric sculpticells visible in lateral depressions. Setal formula ++/++/‒2++.

Male genitalia (n = 1). Aedeagal median lobe moderately broad dorsoventrally, apex narrowly rounded and slightly projected beyond ostium (Fig. 15A); flagellar plate large and lightly sclerotized; right paramere narrow and elongate (Fig. 16A), ventral margin setose, with >20 setae along margin, setae more densely distributed in basal half; left paramere broadly subquadrate basally, apically attenuated into a whip-like apex.

Female reproductive tract (n = 1). Bursa copulatrix broadest at midlength, its surface membranous and covered with pleat-like wrinkles, apex narrowed (Fig. 17A); spermathecal duct narrow, elongate, about twice length of spermathecal reservoir; basal gonocoxite broad, 2–4 setae along apical margin (Fig. 18A), several setae along median margin; apical gonocoxite broadly rounded apically, mesal and lateral margins subparallel; a single broad lateral ensiform seta, its length about 1/3 length of apical gonocoxite; apical nematiform setae in large, apical positioned sensory furrow.

Lectotype female (MCG) designated by Straneo (1941): specimen glued to platen // Lateralis / Cast. / Paroo riv. // Australia / Paroo River / Coll. Castelnau // LECTOTYPUS / Forticosomus / lateralis / Castelnau, 1867 / des. S.L. Straneo, 1941 [orange label] // Forticosomus / lateralis // Mecyclothorax / lateralis Cast / holotypus / Det. B.P. Moore ʽ68 // MUSEO GENOVA / Coll. Castelnau. A female paralectotype accompanies the lectotype. It bears only the newer “Australia / Paroo River / Coll. Castelnau” label and Straneoʼs orange PARALECTOTYPUS label. As noted by Straneo (1941: 89), the paralectotype is teneral. For C. cinctipennis Blackburn, holotype (BMNH; label data not recorded): Torrens River, S.A. (Moore et al. 1987).

Nomenclatural note. In the paragraph within which Chaudoir (1873) combined Phorticosomus minutus Castelnau with Simodontus Chaudoir (pp. 113–114), Chaudoir writes “Je crois que son Ph. lateralis est encore une espèce de Simodontus, qui mʼa semblé différente du curtulus (p. 114).” Such a statement falls short of a nomenclatural act proposing a valid new combination. Csiki (1929) interpreted Chaudoir (1873) to have officially combined Ph. lateralis with Simodontus, however his listing the page of that action as Chaudoir (1873: 113) suggests that he did not see the work personally, thus leading him to give Chaudoir credit for a nomenclatural act that Chaudoir did not commit. Sloane agreed that Chaudoir did not combine Ph. lateralis with Simodontus, writing: “Ph. lateralis is a species of Mecyclothorax. Ph. minutus, from the Paroo River, has been examined by Chaudoir, who referred it to Simodontus ... (Sloane, 1915, 462).” Mooreʼs labelling of the lectotype as holotype was corrected in Moore et al. (1987), where the presence of a paralectotype was also noted.

This species is distributed in interior Victoria, western New South Wales and southeastern South Australia (Fig. 19A). Localities and repositories for non-type specimens I have examined include: SA: Lucindale (MCZ, 1); Pt. Lincoln (MVM, 1); VIC: Bendigo (MVM, 1); Birchip (MVM, 1); Evansford (BMNH, 15); Lake Hattah (MVM, 1); Maldon (MVM, 1); Wedderburn (MVM, 1). These records represent beetles collected between 1911 and 1950, none with any ecological data, and so nothing can be reported with confidence concerning the ecological preference of this species: but see M. minutus below. This species is polymorphic for flight-wing configuration, with 3 brachypterous individuals observed among the 27 beetles examined; 2 of the individuals from Evansford exhibit wings that are broad, slightly more than half as long as the elytra, and without a reflexed apex. Other examined individuals have vestigialized wings, with the wing stubs shorter than the metanotum.

Figure 7. 

Male aedeagus, right view (unless stated otherwise), for Mecyclothorax (Eucyclothorax) spp.: A, M. moorei, NSW: Bellangry For.; B, M. moorei, NSW: Bellangry For.; C, M. punctatus, VIC: Sea Lake; D, M. curtus, VIC: Bendigo; E, M. blackburni, WA: Fremantle; F, M. darlingtoni, QLD: Woondom For. Res.; G. same specimen left view; H, M. darlingtoni, QLD: 30 mi. N Brisbane.

(n = 5). This species shares the rufous to brunneous body color (Fig. 6A–B) and lack of the parascutellar seta with M. lateralis, however the beetles are smaller–standardized body length 4.9–5.7 mm–and the elytral lateral and apical margins do not markedly contrast with the elytral disc. In teneral individuals the margins may appear somewhat paler, partly due to reflection of the underlying abdominal tergites through the cuticle, and partly due to a smoky infuscation of the median elytral disc. However, the difference in coloration from disc to margin is gradual, never marked. Males exhibit 2 setae each side of the apical ventrite margin, for a total of 4 apical setae, differing from M. lateralis which retains the plesiomorphic single seta per side: i.e. a total of 2 apical abdominal setae. The labrum is distinctly emarginate apically, the 2 sides subangulate medially. The ligula is truncate apically, with the 2 apical setae separated by 4 setal diameters, the ligular surface longitudinally depressed between the setal articulations. The paraglossae elongate, extending twice as far beyond the ligular apical margin as the distance from paraglossal base to ligular margin. The pronotum is transverse, with the depressed median base margined at the disc with strigose punctures (Fig. 2K); MPW/BPW = 1.47–1.56, MPW/PL = 1.42–1.51. The prosternal process is broad and only slightly depressed between the coxae, convex anterad the coxae. The prosternal anteapical groove is deep and narrow, smooth at depth. As in M. lateralis, the metepisternum is abbreviated, with lateral length about 2× maximum width, and the metathoracic wings are vestigial in examined material. The suture between visible ventrites 1 and 2 is sinuous, with a broad circular depressed area posterad the sinuosity. Microsculpture of the head is reduced, with frons and vertex glossy, the surface interrupted only by scattered micropunctures. Similarly the pronotal disc and median base are glossy, with indistinct transverse lines visible in places. The elytral surface is glossy with micropunctures covering the surface of the disc, the apex with margins of transverse sculpticells upraised. Ventrally, the thoracic ventrites are glossy; abdominal ventrites glossy medially but with swirling transverse sculpticells visible in lateral depressions. Setal formula ++/++/‒2++.

Male genitalia (n = 1). Aedeagal median lobe moderately broad dorsoventrally, apex narrowly rounded and not projected beyond ostium (Fig. 15B); flagellar plate large and lightly sclerotized; Right paramere narrow, elongate, >20 setae along ventral margin, setae more densely packed in basal half, dorsal margin with ~4 small setae (Fig. 16B); left paramere slightly broadened basally, evenly narrowed to apex.

Female reproductive tract (n = 1). Bursa copulatrix broadest at midlength, its surface membranous, apex narrowed into an elongate projection covered with pleat-like wrinkles (Fig. 17B); spermathecal duct moderately narrow, elongate, about twice length of spermathecal reservoir; basal gonocoxite broad, 2–4 setae along apical margin (Fig. 18B), 1 seta at apicomedial angle, and several setae along median margin; apical gonocoxite broadly rounded apically, mesal and lateral margins subparallel; a single narrow, acuminate lateral ensiform seta, its length about 1/4 length of apical gonocoxite; apical nematiform setae in large, apical positioned sensory furrow.

Figure 8. 

Male parameres of Mecyclothorax (Eucyclothorax) spp., ectal view, right paramere above in each pair, left paramere below: A, M. moorei, NSW: Bellangry For.; B, M. punctatus, VIC: Sea Lake; C, M. curtus, VIC: Bendigo; D, M. blackburni, WA: Fremantle.

For P. minutus, lectotype female (MCG) designated by Straneo (1941): specimen glued onto elongate trapezoidal point // riv. N.H. / Murray [blue label] // minutus / Cast. // N(ova) Hollandia / Riv. Murray / Coll. Castelnau //Forticosomus / minutus / Cast. // LECTOTYPUS / Forticosomus / minutus / Castelnau, 1867 / des. S.L. Straneo, 1941 // Mecyclothorax / minutus Cast / holotypus / Det. B.P. Moore ʽ68 // MUSEO GENOVA / Coll. Castelnau. Mooreʼs labelling of this specimen as holotype, as done for M. lateralis above, is corrected in Moore et al. (1987) to reflect Castelnauʼs lack of mention of how many specimens from Paroo River were before him at description. For C. fortis Blackburn, lectotype male (BMNH) hereby designated: specimen mounted on white card with “913 T” on obverse, black and red lines at base, the red crossed by two triangles // round, red-margined Type label // Blackburn Coll. / 1910-236 // Cyclothorax fortis, Blackb. // Lectotype / Cyclothorax / fortis / Blackburn / J.K. Liebherr 2006 (black-margined red label). Female paralectotype (SAMA): specimen card-mounted with ventral surface upward, card with black and red line, “913” in red ink // S. Australia / Blackburn // Cyclothorax / fortis, Bl / Co-type // Paralectotype ♀ / Cyclothorax / fortis Blackburn / det. J.K. Liebherr 2004 [black margined, red label]. The lectotype (BMNH) was assigned based on its occupation of the first locality, Port Lincoln, mentioned in Blackburnʼs description (Blackburn 1889: 1391), with the paralectotype assigned to the second locality mentioned by Blackburn; “Yorkeʼs Peninsula.” Thus Port Lincoln becomes the type locality for the Blackburn name.

This species exhibits a bicentric distribution, occupying the interiors of Western Australia, and New South Wales, South Australia and Victoria (Fig. 19B). Distributional records and institutional repositories for examined non-type specimens include: NSW: Bogan R. (MCZ, 1); Lake Urana Nat. Res. (EMEC, 17); SA: Koongawa, 4 mi. ESE (ANIC, 1); Pt. Augusta (BPBM, 7); Birchip (MVM, 1); Mallee Dist. (MVM, 3); Nyah (MVM, 1); WA: Burracoppin, 129 mi. S (WAM, 1); Coongardie, 85 km W (BPBM, 3); Hineʼs Hill, 10 mi. SW Merredin (CAS, 4); Hughden Rock (WAM, 2); Mullewa (MCM, 1; MCZ, 10); Newman Rock (ANIC, 1); Ravensthorpe, 50 mi. E (CAS, 1); Southern Cross, 14 mi. W (CAS, 1); Yellowdine, 18 km S (UASM, 2). Beetles constituting the sizable series collected by K.W. Will at Lake Urana Nature Reserve were found under rocks near pools of water in forest, by headlamp at night. As with most carabid beetles living in dry habitats, such nighttime entomological activity is a requisite for obtaining more than the odd specimen. Given the close, adelphotaxon relationship between M. minutus and M. lateralis, this type of habitat is suggested as the situation within which to find M. lateralis as well.

(n = 5). Moore (1984) deferred reliable diagnosis of this species from M. punctipennis based on the configuration of the male aedeagus (e.g. Fig. 15C–D), however the pronotal lateral margins are reliably narrower in individuals of this species (Fig. 2L). Although variable melanization of the pronotal disc and marginal depressions may confuse the appearance of the width of the lateral depressions, other aspects of the pronotum may be used, including: 1, punctation of the median base, with about 20–22 punctures each side in this species versus nearly 30 deeper punctures each side in M. punctipennis (Fig. 2L–M); 2, median base juncture with pronotal disc lined with deep, nearly round to moderately elongate punctures in this species, versus distinctly strigose depressions with 1–2 punctures in each depression for M. punctipennis. Microsculpture also varies unambiguously between the two species: 1, in M. ambiguus a transverse mesh visible over portions of the pronotal disc and median base, best viewed adjacent to areas of reflected microscope light, versus glossy pronotal disc and median base, with only indistinct transverse lines within irregularities of the cuticular surface in M. punctipennis; and 2, discal elytral intervals with well-developed transverse mesh, sculpticell breadth 2–3× length, and surface subiridescent in M. ambiguus, versus discal elytral intervals glossy, with at most indistinct transverse sculpticells visible at margins of fields of reflected light, the surface without any iridescence in M. punctipennis. The depth of elytral striae varies on the elytral apex, however in M. ambiguus, interval 8 is broadly convex adjacent to the well-developed stria 7 between the subapical and apical elytral setae, whereas in M. punctipennis, interval 8 is internally subcarinate and angularly depressed laterally (Liebherr 2012a, fig. 7). For M. ambiguus, the mentum tooth has sides obtuse, the apex broadly rounded, whereas M. punctipennis is characterized by an acute mentum tooth with apex tightly rounded. The prosternal anteapical groove is deep and smooth here, but broader and distinctly punctate laterally, though smoother and slightly irregular medially, in M. punctipennis. In addition, the depressions surrounding the dorsal elytral setae span 1/4–1/2 of elytral interval 3 in this species, but up to 1/2–3/4 of the interval width in M. punctipennis. Standardized body length 5.0–5.7 mm. Setal formula ++/++/+2++.

Male genitalia (n = 3). Aedeagal median lobe gracile, narrow dorsoventrally relative to length, apex broad, expanded both ventrally and dorsally resulting in a nearly straight apical face (Fig. 15C); ostial ventroapical operculum well developed as a broadly triangular sclerite; flagellar plate very large, well-sclerotized with longitudinal ridges; aedeagal internal sac bearing a ventral spicular sclerite; right paramere slightly broadened basally, evenly narrowed to apex, ventral surface with ~20 setae along margin, additional very small setae may be present near apex (Fig. 16C); left paramere slightly expanded in basal half, apically narrowed to whip-like apex.

Female reproductive tract (n = 2). Bursa copulatrix elongate, columnar, length about 3× diameter when pressed under cover slip, surface membranous, translucent, wrinkled (Fig. 17C); spermathecal duct entering bursa copulatrix mediodorsally, duct length about 2× length of spermathecal reservoir; spermathecal gland duct long, >3× length of spermathecal reservoir; basal gonocoxite apical margin with 4 setae, 1 setae at apicomedial angle, and several smaller setae along medial margin (Fig. 18C); apical gonocoxite broad basally with 2 short, stout lateral ensiform setae, apex narrowly rounded; apical nematiform setae in sensory furrow near apex of apical gonocoxite.

Dissected and pinned male Lectotype (ZMHU): 3294 // ring sclerite and aedeagus on card // ambiguus / Er. / Van Diemens Land / Schayer [blue label] // LECTOYPE (red label) Mecyclothorax / “Anchomenus” / ambiguus / Erichson 1842 / designated by / B. P. Moore. An additional two male and one female paralectotypes (ZMUH) accompany the lectotype.

Figure 9. 

Female reproductive tract and gonocoxae of Mecyclothorax (Eucyclothorax) spp., ventral view: A, M. moorei, NSW: Werrikimbe N. P.; B, M. punctatus, VIC: Birchip; C, M. curtus, SA: Manangatan; D, M. blackburni, WA: Harvey; E, M. darlingtoni. QLD: Woondom For. Res; F, M. lophoides, ACT: Smoker’s Gap; G, M. eyrensis, SA: Telowie Gorge; H, M. peryphoides, ACT: Black Mountain; I, M. cordicollis, NSW: Gosford.

This species is distributed throughout southeastern Australia including Tasmania and King Island (Fig. 19C). Localities and repositories for examined material include: NSW: Bodalla (MCZ, 4); Braidwood (MCZ, 41); Breakfast Ck. (MVM, 1); Cabramatta (BMNH, 1); Comboyne plateau (MCZ, 4); Cumberland (BMNH, 1); Greta (AMNH, 1); Kosciuszko N. P. (CUIC, 8; MCZ, 7); Mt. Wilson (FMNH, 1); Narrabeen Lagoon (FMNH, 5); Richmond R. vic. Wiangaree (MCZ, 8); Tallaganda St. For. (CAS, 1); Uralla (CAS, 3); SA: Adelaide (CAS, 1); Adelaide, 18 km E, Carey, 3 km SE (FMNH, 1); Adelaide, 8 km S, Waterfall Gully (FMNH, 1); Blackwood, Sturt Gorge Res. (ZMUC, 13); Goose Island (MVM, 3); Norton Summit, 10 km E Adelaide (FMNH, 2); Williamstown (FMNH, 1); Yorketown (CAS, 2; FMNH, 13); TAS: Burnie (MCZ, 8); Cockle Ck. (MCZ, 2); Florentine R. (MCZ, 3); Goulds Country (CMNH, 2); Hartz N. P. (MCZ, 2); Hastings (MCZ, 3); Hobart (MVM, 3); King Is. (MVM, 2); Launceton (FMNH, 1; MCZ, 1); Mersey R. Vy. (MCZ, 2); Mt. Ben Lomond (MCZ, 6); Mt. Field, base (MCZ, 7); Mt. Wellington (MCZ, 1); Parratah (MVM, 2); Queenstown (MCZ, 16); Smithton (CAS, 7); Tarraleah (MCZ, 3); Waldheim (MCZ, 1); Zeehan (CAS, 1); VIC: Ballarat (BMNH, 2); Bogong (MVM, 2); Dandenong (CAS, 2); Frankston (MVM, 1); Ferntree Gully (MVM, 1); Lakes Entrance (MVM, 1); Mt. Buller (MCZ, 1); Mt. Donna Buang (MCZ, 15); Mt. Hotham (MCZ, 3); Mt. Wogwog (EMEC, 1); Oakleigh (MVM, 2); Olinda (BMNH, 2); Penshurst (CUIC, 7); Port Melbourne (MVM, 2); Portland to Pt. Fairy (MCZ, 2); Pretty Vy. Dam (MVM, 1); Warburton (MVM, 6); Warrendyte (MVM, 1); Whiskey Ck. (MVM, 3); Yarra Glen (MVM, 1).

(n = 5). For purposes of this review, all diagnostic external characters that distinguish this species from M. ambiguus–and therefore all other Australian species–are presented under M. ambiguus. Standardized body length 5.0–5.8 mm. Setal formula ++/++/+2++.

Male genitalia (n = 3). Aedeagal median lobe gracile, narrow dorsoventrally relative to length, the apex well extended beyond ostium, the tip downturned (Fig. 15D); ostial ventroapical operculum well developed, an elongate triangular sclerite; flagellar plate large, bearing longitudinal sclerotic ridges (Fig. 15E); base of aedeagal internal sac bearing a ventral spicular sclerite; right paramere narrow, elongate, bearing >12 setae along the ventral margin, 4 small setae long dorsal margin (Fig. 16D); left paramere narrow basally, narrowed to elongate, attenuated whip-like apex.

Female reproductive tract (n = 2). Bursa copulatrix elongate, columnar, length about 2× diameter when pressed under cover slip, surface thickened, wrinkled, (Fig. 17D); spermathecal duct entering bursa copulatrix mediodorsally, duct length about 2× length of spermathecal reservoir; spermathecal gland duct long, ~1.5× length of spermathecal reservoir; basal gonocoxite apical margin with 3 setae, 1 seta at apicomedial angle, and several smaller setae along medial margin (Fig. 18D); apical gonocoxite broad basally with 2 acuminate lateral ensiform setae, apex acuminate; apical nematiform setae in subbasal sensory furrow.

For M. punctipennis, lectotype male (ANIC): Gayndah, Queensland (Moore 1984). For C. obsoletus Blackburn, two syntypes (SAMA): Port Lincoln, S.A. (Moore 1984, Moore et al. 1987).

Figure 10. 

Left gonocoxa of Mecyclothorax (Eucyclothorax) spp., ventral view: A, M. moorei, NSW: Werrikimbe N. P.; B, M. punctatus, VIC: Birchip; C, M. curtus, SA: Manangatan; D, M. blackburni, WA: Harvey; E, M. darlingtoni. QLD: Woondom For. Res; F, M. lophoides, ACT: Smoker’s Gap; G, M. eyrensis, SA: Telowie Gorge; H, M. peryphoides, SA: Blackwood; I, M. cordicollis, NSW: Gosford.

This species is broadly distributed in numerous habitats across Australia (Fig. 19D). Recorded collection localities range in elevation from sea level to over 2000 m near the summit of Mt. Kosciuszko. These beetles are at home in leaf litter on the floor of Eucalyptus forests, under dense mats of dead leaves surrounding the bases of Xantharrhoea (Asphodelaceae) grass trees, in tussock grass clumps of high-elevation grasslands, under wrack on sea beaches, and in home gardens in urban settings. The species is monomorphically macropterous, with adults often collected at lights in great profusion.

Even given this speciesʼ catholic ecological preferences and propensity for winged flight, its geographic distribution is discontinuous across Australia (Fig. 19D). Moreover, the Western Australian populations of this bicentric species distribution interact little if at all with coastal populations east of the Great Australian Bight, based on geographic restriction of polymorphic male genitalic chiral antisymmetry to the populations inhabiting Western Australia (Liebherr and Will 2015). In contrast, all eastern populations monomorphically comprise males with plesiomorphic genitalic torsion, whereby the right side of the aedeagus is held ventrally when in repose. The Western Australian populations vary greatly in the proportions of left- and right-torsioned males, demonstrating that their mutual geographic isolation is great enough to preclude extensive homogenizing dispersal among populations.

Baehr (2000) reported a 1998 record for M. punctipennis from Rocky Cape N. P. as the first Tasmania record. However, Darlington material (MCZ) indicates M. punctipennis was present at Hobart in 1956–1957 (Liebherr and Will 2015). Tasmanian localities and repositories represented in material examined for this review (Fig. 19D) include: Corinna, West Tasmania (MCZ, 4); Hobart (MCZ, 8); Great Lake, north end (MCZ, 5); Waldheim nr. Cradle Mtn. (MCZ, 1); Zeehan, north (MCZ, 3).

Figure 11. 

Distributional ranges of 11 Mecyclothorax spp. assigned to subgenus Eucyclothorax, plus newly described subspecies of M. lewisensis.