Theprisa darlingtoni Liebherr & Porch, 2021

Liebherr, James K., Porch, Nick, Shaw, Matthew, Sinclair, Bronte E. & Maddison, David R., 2021, Systematic revision of the trans-Bassian moriomorphine genus Theprisa Moore (Coleoptera, Carabidae), ZooKeys 1044, pp. 339-373 : 339

Theprisa darlingtoni Liebherr & Porch sp. nov. Figures 2B , 3E , 4E , 5E , 6E , 8E , 10 , 11

Holotype

♂ (ANIC deposition of MCZ specimen): N. of Zeehan / Jan. '57 Tas / Darlingtons // MCZ ENT 00731903 // HOLOTYPE ♂ / Theprisa / darlingtoni / J.K. Liebherr & N. Porch 2020 (red-margined black label). Field data for the type locality include: low, wet sclerophyll forest, rain forest in places (but much cleared); Jan 17-19, 1957; usual ground methods (under logs or stones, alongside water, or drowning leaf litter in water); mostly circa 8 miles north, and along Rosebery Rd. ( Darlington 1960: 125).

Allotypic paratype

♀. Same label and deposition as holotype.

Paratypes.

Same label as holotype (MCZ, 8); Tasmania: Mt. Ben Lomond [sclerophyll forest, wet gullies; logs, stones, waterside, drowning, under bark], 900-1200 m, [5-10]-iii-1957, Darlingtons(MCZ, 1); Corinna [temperate rain forest incl. Nothofagus ; logs, stones, waterside, drowning; low elevation], [14-17]-iii-1957, Darlingtons (MCZ, 5); Whyte R., Corinna Rd. [sclerophyll forest, temperate rain forest incl. Nothofagus ; logs, stones, waterside, drowning; low elevation], [17-18]-iii-1957, Darlingtons (MCZ, 4); Blue Tier, NE Tasmania [temperate rain forest incl. Nothofagus , partly cleared; logs, stones, waterside, much drowning debris], ~ 600 m, [26-31]-iii-1957, Darlingtons (MCZ, 1); Gordon R. Gorge, btw. Sir John Falls & Pyramid Is., 42.5749°S, 145.7169°E, 3-ii-1979, Howard (TMAG, 1); Frankland Range, 42.92°S, 146.01°E, 1-i-1991, Mesibov (TMAG, 1); Corinna, NW Tasmania, 4-xii-1999, Bouffard (CMNH, 1).

Diagnosis

(n = 5). T. darlingtoni is most similar to T. montana based on large body size; 7.4-8.0 mm versus 7.0-8.9 mm for T. montana . In both species the elytra are broad basally, here HuW/MEW = 0.66-0.67, with the lateral margins subparallel (Figs 1C , 2B ), and the elytral striae smooth. But T. darlingtoni exhibits a much more pronounced tubercle laterad the deep, arcuate depression bordering the pronotal median base, and the pronotal lateral margins are only briefly sinuate laterad the basal pronotal seta (Fig. 2B ), not concavely curved laterad the entire laterobasal depression as in T. montana (Fig. 1C ). The elytral lateral marginal depression is also much broader in T. darlingtoni , with the rufous coloration of the depression contrasted to the rufopiceous interval 8, whereas the narrow lateral depression of T. montana is only slightly paler than interval 8. Cuticular microsculpture is well developed, with the vertex covered by a shallow but evident transverse mesh, the pronotal disc bearing a shallow transverse mesh, sculpticell breadth 4 × length, and the elytral surface with silvery subiridescence due to the presence of dense transverse lines across the convex elytral intervals. The apical abdominal ventrite of both male and female bear three seta each side along the margin, the median pair in line with the outer two bilateral pairs.

Description.

Head robust, ocular lobe protruded, its juncture with gena obtuse; eye diameter small relative to elongate head capsule, surface not projected beyond curve of ocular lobe, EyL/OLL = 0.69-0.71, but ocular ratio moderate, 1.40-1.46, and horizontal line across eye bisecting 21-25 ommatidia; antennal scape robust, diameter 1.4 × apical diameter of pedicel; antennomeres 2 and 3 glabrous except for one dorsal seta on the pedicel and apical ring of setae on 3; antennae elongate, antennomere 9 maximal breadth 2.14 × length; frontal groove broad, deep, the surface of impression irregularly strigose, the two grooves laterally arcuate, defining an ovoid raised area on frons, divergent from frontoclypeal suture to lateral margin of clypeus; broadly, deeply emarginate apical margin of labrum lined with six (rarely seven) setae, six smaller setae visible along anterolateral labral margin; mentum tooth narrowly rounded apically, sides acutely divergent; maxillary stipes trisetose, the three setae on the base in either a triangle with apex upward, or in an irregular horizontal line; ligula truncate apically, narrowed basally, trumpet shaped, its two apical setae separated by three setal diameters; paraglossae elongate, total length 3 × distance from paraglossal base to ligular apical margin. Pronotum moderately transverse, lateral margins straight to slightly concave anterad acutely projected margin at basal pronotal seta; basal margin smoothly, convexly curved across width, the unmargined median base not projected beyond curve defined by distinct lateral margins posterad laterobasal depressions; median base depressed below disc, smooth basally, an arcuate line of isolated punctures each side along juncture with disc; longitudinal depression consisting of medial, deep, arcuate impression and a nearly smooth lateral tubercle, a few small punctures laterad tubercle near lateral marginal depression inside hind angles; median longitudinal impression present on median base as a series of isolated punctures, on disc consisting of lenticular depression at median base-disc juncture, and an irregular, deeply incised impression on disc; anterior transverse impression obsolete, very shallow, the median longitudinal impression continued anterad halfway across flat anterior collar of pronotum; pronotal anterior margin smooth medially, a narrow marginal bead increasingly more well developed in outer 2/3 of width each side; front angles protruded, tightly rounded; lateral marginal depression moderately narrow, broad enough so that cuticular sculpticells visible mesad beaded, raised margin; margin of lateral pronotal seta articulatory socket adjoining lateral marginal depression. Prosternum deeply canaliculate from prosternal process halfway to anterior prosternal margin, smooth over much of surface but with a few punctures or strigae at proepisternal suture anterad coxal cavity; proepisternum smooth, sutural groove with proepimeron smooth and deep. Elytra broadly ellipsoid, MEW/EL = 0.75-0.78, moderately convex, sides meeting lateral marginal depression nearly vertically, disc between striae 4 flat at midlength; basal groove arcuate, pitted at bases of striae 1-5, a small acute tooth present at juncture of groove and lateral marginal depression; stria 8 very deep and continuous between anterior and posterior series of lateral setae; apical carina of interval 8 narrowly upraised along stria 7, interval 8 a vertical lateral carina there; subapical sinuation abrupt, the internal plica visible ventrad deepest part of sinuation. Mesepisternum broadly punctate, ~ 19-22 punctures in 3-4 confused vertical rows. Metepisternum trapezoidal, maximum width subequal to lateral length; metepisternal-metepimeral suture complete. Legs gracile, the femora elongate and meso- and metatibiae only slightly broadened in apical half; metatarsomere 1 moderately elongate, length 0.20 × tibial length, lateral sulci present on mesal and lateral faces just dorsad the ventrolateral setae. Abdominal ventrites 2-6 smooth laterally, but hind margins of ventrites 2-5 concavely sinuate laterally, the sutures deeper in association with sinuation; suture between ventrites 1 and 2 deep, slightly curved anteriorly at midlength; suture between ventrites 2 and 3 complete laterally.

Male genitalia (n = 5). Aedeagal median lobe elongate, moderately robust, base broadly open on right side, basal margin heavily sclerotized dorsad basal opening (Fig. 3E ); apical face joined to slightly arcuate ventral margin at right angle, flattened tip not extended beyond ostium, lateral surfaces of apex smooth or with only several indistinct pits; internal sac largely membranous, with only very elongate flagellum and associated basal articulatory sclerites darker (Figs 3E , 4E ); right paramere elongate, slightly angled at midlength, apex narrowly rounded, slightly asymmetrical in outside lateral view (Fig. 5E ), bearing 10-19 short and long setae along ventral margin in apical half, 2-4 setae on dorsal surface near apex, two or three setae at apex; left paramere broadest in basal half, narrowed in apical half but broadened apically to broadly rounded tip, ventral margin with 3-7 setae near apex, dorsal surface glabrous or with a single seta, and apex with two or three longer setae; antecostal apodeme of abdominal segment IX angulate distally, lateral arms gracile, their distal juncture only slightly broadened (Fig. 6E ).

Female reproductive tract (n = 1). Bursa copulatrix columnar, length 1.5 × maximum breadth compressed under microslide cover slip, vagina translucent, apical portion of bursa staining more darkly with Chlorazol black (Fig. 11 ); helminthoid sclerite present, broadly rounded apically, not extended beyond juncture with spermathecal duct; spermathecal duct thick, sclerotized, coiled sinuously to spermatheca, length twice that of annulated spermathecal reservoir; spermathecal gland duct very thin, length half that of spermathecal reservoir which it joins at reservoir base; spermathecal gland comprising sclerotized stem plus membranous reservoir bearing numerous ductules; gonocoxa bipartite, basal gonocoxite 1 with single apical fringe seta, median surface glabrous, membranous ramus present (Fig. 8E ); apical gonocoxite 2 with base extended laterally, lateral margin arcuate, apex narrowly rounded; two lateral ensiform setae and one dorsal ensiform seta present; two apical nematiform setae set in fossa in apical 1/4 of apical gonocoxite length.

Etymology.

This species honors Professor Philip J. Darlington, Jr., who among his many roles, systematically collected carabid beetles across Australia, and served as the first post-doctoral supervisor for Dr. Terry L. Erwin ( Kavanaugh 2020). Terry championed a Darlingtonian world view regarding the diversification of carabid beetles ( Erwin 1981), never losing sight of those questions that tied both men to primary observations made over long careers spent in the field.

Distribution and habitat.

Theprisa darlingtoni is known from localities spanning northern and western Tasmania, from Blue Tier on the east to Corinna in the northwest, to the Frankland Range in southwest Tasmania (Fig. 10 ). Collecting localities lie within the Ben Lomond, Central Highlands, and West biogeographic regions (IBRA 2004). It has been collected syntopically with T. convexa at three localities: Corinna (Bouffard, CMNH), Whyte River at Corinna Road (Darlingtons, MCZ), and Zeehan (Simson, SAMA, and Darlingtons, MCZ), although these respective 1891 and 1957 collections were separated by 66 years. The five sites at which the Darlington family collected this species (Blue Tier, Corinna and nearby Whyte River, Mt. Ben Lomond, and Zeehan) were reported to support sclerophyll forest, wet sclerophyll forest, temperate rain forest with Nothofagus , and wet gullies, with specimens at all localities collected under logs and stones, along water sources, and via immersion of leaf litter in standing water ( Darlington 1960).

Phylogenetic analysis

Parsimony analysis finds two cladograms of 380 steps, with the strict consensus collapsing one node within representatives of Neonomius (Fig. 12 ). Monophyly of the five Theprisa spp. is recovered and this clade is adelphotaxon to a clade comprising Spherita and Sitaphe . This sister-group relationship was not recovered in Liebherr (2020), but that analysis was comprehensive taxonomically, and this less so as it was focused only on demonstrating monophyly of Theprisa . In the current analysis, there is also very high support for the adelphotaxon relationship of Sitaphe + Spherita (Fig. 12 ). There is substantial support within the morphological character data for Theprisa monophyly, with the genus obtaining a decay index value of 6. Several characters unambiguously support Theprisa monophyly. The uniquely pitted aedeagal apex observed in four of the five species (character 97, state 1; Fig. 3A-D ) has not been observed in any other moriomorphine. The prosternum is grooved laterally about halfway along its length each side near the proepisternum (character 41, state 1), a condition not observed in the included taxa except for New Zealand's, Tarastethus spp. and Trichopsida pretiosa (Broun), though those taxa exhibit a series of pits in this position. The metepisternum and metepimeron are fused laterally (character 64, state 1), no doubt in association with the extremely reduced metathoracic flight-wing apparatus. And the eyes are small, with the ocular lobe ratio (EyL/OLL) less than 0.72 (character 18, state 0), though T. montana exhibits variation in this character with some individuals having larger eyes. Finally, the spermathecal gland has a well-sclerotized stem (Figs 7 , 11 ; character 103, state 1), a condition evolved independently in the clade subtended by Pterogmus (Fig. 12 ; Liebherr 2019).

Figures

Figure 1. Dorsal habitus illustrations of Theprisa spp.; range of standardized body lengths indicated A T. otway female B T. convexa female C T. montana female.

Figure 2. Dorsal habitus illustrations of Theprisa spp.; range of standardized body lengths indicated A T. australis male B T. darlingtoni, female.

Figure 3. Male aedeagus including parameres of Theprisa spp., right, or anatomically ventral view A T. otway B T. convexa C T. montana D T. australis E T. darlingtoni. Abbreviations: ad, apical divot; af, apical face; fl, flagellum; lp, left paramere; mf, microtrichial field; rp, right paramere.

Figure 4. Male aedeagus internal sac in everted condition for Theprisa spp. A T. otway, right view B T. convexa C T. montana D T. australis E T. darlingtoni. Abbreviations: fl, flagellum; mf, microtrichial field; rp, right paramere. Scale bars: 0.5 mm

Figure 5. Parameres of male aedeagus for Theprisa spp., outside lateral view. Right paramere shown above left paramere A T. otway B T. convexa C T. montana D T. australis E T. darlingtoni. Abbreviations: lp, left paramere; rp, right paramere.

Figure 6. Structures associated with abdominal segments VIII and IX for Theprisa spp., dorsal view A T. otway B T. convexa C T. montana D T. australis E T. darlingtoni. Abbreviations: VIII, antecostal apodeme of abdominal segment VIII; IX, antecostal apodeme of abdominal segment IX; sp, spiracle of segment VIII; t, tergite of segment IX. Scale bars: 0.5 mm.

Figure 7. Female reproductive tract of Theprisa spp., ventral view A T. otway B T. convexa C T. montana D T. australis. Abbreviations: bc, bursa copulatrix; co, common oviduct; dgd, defensive gland efferent duct; gc, gonocoxa; hg, hindgut; hs, helminthoid sclerite; sd, spermathecal duct; sg, spermathecal gland; sgd, spermathecal gland duct; sgs, spermathecal gland stem; sp, spermatheca; v, vagina. Scale bars: 0.50 mm.

Figure 8. Left gonocoxa, ventral view of Theprisa spp., ventral view A T. otway B T. convexa C T. montana D T. australis E T. darlingtoni. Abbreviations: afs, apical fringe seta (e) of gonocoxite 1; ans, apical nematiform setae; des, dorsal ensiform seta; gc 1, basal gonocoxite 1; gc 2, apical gonocoxite 2; les, lateral ensiform setae; r, ramus.

Figure 10. Distributional records for Theprisa spp.: T. australis (circle);  T. darlingtoni (square).

Figure 11. Female reproductive tract of Theprisa darlingtoni, ventral view; scale bar = 0.50 mm. Abbreviations: bc, bursa copulatrix; co, common oviduct; gc, gonocoxa; hg, hindgut; hs, helminthoid sclerite; sd, spermathecal duct; sg, spermathecal gland; sgd, spermathecal gland duct; sgs, spermathecal gland stem; sp, spermatheca; v, vagina.

Figure 12. Strict consensus of two equally parsimonious, 380 - step cladograms including Theprisa spp. and cladistically neighboring moriomorphine taxa of the subtribe Tropopterina (see text); consensus cladogram length 382 steps, CI = 0.43, RI = 0.66. Character numbers are shown above cladogram edges, character states below. Cladogram root placed so tree topology is compatible with the more inclusive cladogram of Liebherr (2020: fig. 1). Filled squares represent unique state transformations on cladogram, open squares indicate state transformations that occur more than once on cladogram. Clades of this analysis previously recovered in Liebherr (2020) are indicated by pointers numbered 1; the clade subtended by Pterogmus reported in Liebherr (2019) indicated by the pointer numbered 2. Decay indices (i. e., Bremer Support values) shown beneath cladogram edges for all internal edges of the cladogram. Geographic distributions of species are indicated by abbreviations following species epithets: EOZ, southeastern Australia; i. e., A. C. T., New South Wales and Victoria; NNZ, North Island, New Zealand; NZ, New Zealand; SNZ, South Island, New Zealand; QOZ, Queensland; Tas, Tasmania; WOZ, Western Australia