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Systematic revision of the trans-Bassian moriomorphine genus Theprisa Moore (Coleoptera, Carabidae)

Abstract The Australian genus Theprisa Moore, 1963, is taxonomically revised to comprise five species, two newly described: Theprisadarlingtoni Liebherr & Porch, sp. nov. of Tasmania, and Theprisaotway Liebherr, Porch & Maddison, sp. nov. from the Otway Ranges, Victoria. Two previously described species, T.australis (Castelnau) and T.montana (Castelnau), are distributed in the mountains of Victoria. The third previously described species, T.convexa (Sloane) is found in Tasmania. A lectotype is designated for T.convexa because the various syntypes are ambiguously labelled. Cladistic analysis based on morphological characters establishes monophyly of Theprisa relative to the Australian genera Sitaphe Moore and Spherita Liebherr. This and a second clade of Australian genera (Pterogmus Sloane, Thayerella Baehr, and Neonomius Moore) do not form a natural group, but are cladistically interdigitated among two monophyletic New Zealand lineages (Tarastethus Sharp, and Trichopsida Larochelle and Larivière) suggesting substantial trans-Tasman diversification among these groups. Hypothesized relationships within Theprisa are consistent with two bouts of speciation involving the Bass Strait; an initial event establishing T.convexa as adelphotaxon to the other four species, and a more recent event establishing the sister species T.darlingtoni and T.montana. Geographic restriction of T.otway to the Otway Ranges is paralleled by Otway endemics in several other carabid beetle genera, as well as by endemics in numerous other terrestrial arthropod taxa. Whereas these numerous Otway endemics support the distinctive nature of the Otway Range fauna, their biogeographic relationships are extremely varied, illustrating that the Otways have accrued their distinctive biodiversity via various means.


Introduction
This revision was initially conceived in a world unlike the one we currently inhabit. As the 2019 coronavirus pandemic spread worldwide, workplaces were shuttered, air travel and transportation changed to essential deliveries, and all of us physically distanced to preserve our collective health. It was also the time that we lost Terry Erwin, taking away a major contributor to the study of biodiversity worldwide (Kavanaugh 2020). A constant theme of Terry Erwin's research was his inevitable response to challenges and adversity. He attacked the challenges directly, taking action to mitigate any negative outcomes, and succeeded by dint of effort and organization. This group of authors took that to heart, and though we are far flung geographically, we assembled a team using the internet to share scientific information, allowing us to remotely assemble the necessary components of this revision. Insect collection curators are charged with both the maintenance and protection of previously described specimens, but also with the development of institutional collections, achieved through use of specimens in support of novel descriptive taxonomy. This activity is increasingly tied to electronic access of collection-based information, yet that information must be of the highest quality to maximally promote truth in science. This contribution brings together information from specimens described during the past 150 years, with those specimens housed in Europe, North America, and Australia. Given the current lockdown of society, only an internet-based effort could produce such a taxonomic product.
The genus Theprisa Moore, 1963, was described as a distinct taxon based on characters of the mouthparts and male genitalia (Moore 1963b), yet the taxonomic unit was first recognized informally by Sloane (1920). In a recent cladistic analysis of Moriomorphini Sloane, 1890, the tribe that includes Theprisa, monophyly of the genus was proposed but only based on inclusion of two Theprisa species (Liebherr 2020). This revision incorporates cladistic analysis including the three previously described species of Theprisa, two newly recognized species, as well as representatives of closely related genera. This allows testing monophyly of Theprisa and proposing its adelphotaxon. Our geographically diverse team, united by the internet, was able to assemble and share data on all available specimens, including all type specimens held in Australian museums. We dedicate the product of our worldwide collaboration to the memory of Dr. Terry L. Erwin. were relaxed for 2 hrs in near-boiling distilled water including several drops of Kodak Photo-Flo detergent (Rochester, NY). For male specimens the invaginated abdominal segments VIII and IX and the included aedeagus were removed using minuten pins, cleared in cold 10% KOH overnight, neutralized in 10% acetic acid, and examined after 24 hours in glycerin. Female dissections involved removal of the entire abdomen, clearing overnight in cold 10% KOH and neutralization in acetic acid, with isolation of gonocoxae and the associated female reproductive tract and hind gut accomplished using fine forceps and minutens. The reproductive tract was stained for 15-30 mins in Kodak Chlorazol Black mixed into methyl cellosolve, with the stained preparation placed into glycerine for 24 hours prior to continued dissection.
Photographs of whole specimens and dissected genitalia and reproductive tracts were made using a Microptics macrophotographic apparatus, mounting a Nikon D1 camera and Microptics K2 lens system, with lighting provided by a three wand fiberoptic photographic strobe unit allowing either direct lighting or transmitted light on a microscope slide stage. Images were captured and stacked using Compose Z5 software (Hadley 2006).

Morphological characters
For diagnostic purposes, various ratios of body dimensions are useful, including those of the head, pronotum, and elytra. Eye development is characterized by the ocular ratio, or maximal head width across the compound eyes divided by the minimal frons width between the eyes. The eyes are borne on an ocular lobe, with the amount of that lobe covered by the eyes quantified as the ocular lobe ratio (EyL/OLL), or the length of the eye measured in dorsal view divided by the distance from the front of the eye to the juncture of the ocular lobe and gena behind the eye. Eye convexity is quantified as the length of the eye in dorsal view divided by depth of the eye (EyL/EyD), that depth measured with the eye's, dorsal margin oriented uppermost in the field of view. Pronotal shape can be used to diagnose species, with MPW/PL, or maximal pronotal width divided by median pronotal length assessing the relative length of the pronotum, and MPW/BPW, or maximal pronotal width divided by basal pronotal width between the hind angles assessing the basal constriction of the pronotum. Elytral configuration is assessed using HuW/MEW, or the distance between the humeral angles divided by maximal elytral width, and MEW/EL, or maximal elytral width divided by elytral length, or the distance from the basal declivity of the scutellum to the apex of the longer elytron just laterad the suture. Standardized body length is quantified as the sum of the distance from the anterior margin of the labrum to the cervical ridge at the back of the vertex, plus PL and EL defined above. For all measurements, the number of individuals measured to estimate variation is indicated parenthetically at the start of the diagnosis.
Cuticular microsculpture can be used to assist diagnosis, with terms for the shapes of sculpticells following Lindroth (1974) and Allen and Ball (1980). Male genitalic armature includes a flagellum on the internal sac (Maddison 1993), with the configuration of that structure providing diagnostic information in both the uneverted and everted condition. Presentation of female gonocoxal setation follows Ball and Shpeley (1983), and female reproductive tract characters, Liebherr and Will (1998). The numbers of individuals dissected to determine genitalic characters is indicated parenthetically at the start of each male genitalia and female reproductive tract section of the descriptions.
Full descriptions of both external and internal genitalic and productive tract characters are presented for both new species. For the previously described species, an extended diagnosis based on external characters is complemented by a full description of the genitalic and reproductive tract characters.

Phylogenetic analysis
Liebherr (2020) presented a morphologically based cladistic analysis of the Moriomorphini, establishing monophyly for genera comprising the tribe. Theprisa was placed in that analysis as a member genus of subtribe Tropopterina, with the two generic representative species placed along a stem of the tree amongst representatives of other genera from Australian and New Zealand (Liebherr 2020: fig. 1). Liebherr (2019), based on personal communication from D. R. Maddison, assessed the inclusion of Thayerella Baehr (2016) in Moriomorphini, concluding that this taxon is a member of a clade also including Pterogmus and Neonomius. Given that the goal of this contribution was revision of Theprisa, the question of Theprisa monophyly gains importance as this paper recognizes five species, two new, to expand the representation of species in Liebherr (2020). As such, phylogenetic analysis focused on Theprisa and its cladistic neighbors is presented to allow assessment of generic monophyly based on the five currently recognized species of Theprisa. The 16 taxa chosen to accompany the species of Theprisa in this analysis included representatives of the following genera: Molopsida White, Tarastethus Sharp, Trichopsida Larochelle and Larivière, Pterogmus Sloane, Neonomius Moore, Spherita Liebherr, and Sitaphe Moore. These genera bracket Theprisa by two cladogram nodes in either direction from the attachment point of Theprisa to the Hennigian comb defining mid-grade members of the subtribe Tropopterina (Liebherr 2020: fig. 1). Taxa sampled from these genera were also restricted to those represented by both male and female specimens. Removing these taxa represented partially by missing data eliminated topological ambiguity from the analysis. By this taxonomic sampling, the goal of testing Theprisa monophyly was undertaken within a reasonable sampling of putatively related taxa, based on complete morphological data for the representative taxa.
The present cladistic analysis is based on 110 characters; 105 ordered binary or multistate, and five unordered multistate characters. The matrix was constructed by culling taxa and associated uninformative characters from the data matrix of Liebherr (2020), while adding character information that includes several relevant new characters for the three additional Theprisa spp. plus Thayerella newtoni Baehr. Charac-ter information was managed using Winclada (Nixon 2002), with analyses run using NONA (Goloboff 1999) and the parsimony ratchet (Nixon 1999(Nixon ) undergoing 200, 1000, and 10,000 iterations. All character and taxon data can be viewed in Suppl. material 2; i.e., the Winclada file with resultant trees derived from the 10,000-iteration ratchet run. Though identical results were obtained under all numbers of iterations of the ratchet, the parsimony results were checked and found identical using TNT (Goloboff et al. 2008;Goloboff and Catalano 2016), running a new technology sectorial search, with ratchet, drift, and tree fusing options; initial addseqs. = 10 and random seed = 1. The matrix was also analyzed with PAUP (Swofford 2002), finding the identical set of parsimonious trees, while also generating decay indices (Bremer 1994) for all internal branches of the cladogram.

Taxonomic treatment
Theprisa Moore, 1963 Theprisa Moore, 1963b: 285 (type species Phersita convexa Sloane, 1920 by original designation. Diagnosis. This genus is diagnosable within Moriomorphini by: 1, mesosternum broad between mesocoxae; 2, mandibular scrobe present, margined ventrally by a lateral expansion; 3, mesotibia gracile, not expanded apically; 4, apical two maxillary palpomeres and apical labial palpomere apparently glabrous; 5, prosternal process not margined apicoventrally; 6, elytral striation nearly complete, striae 1-7 all developed apically, and striae 6-8 as deep or only slightly shallower than striae 1-5 near elytral midlength (Liebherr 2020). The mandibles are moderately elongate, with the mandibular length measured from the anterior condyle 1.7-1.9× distance from anterior condyle to apicolateral margin of labrum. The ligula is broadly rounded to truncate apically, bisetose, with the two setae separated by 2-3 diameters of the setal articulatory sockets. The paraglossae are elongate and thin, their overall length 2-3× the distance from the paraglossal base to the apical ligular margin. The mentum tooth is present, its apex narrowly rounded to subacuminate and its lateral margins upraised. Both lateral and basal pronotal setae are present. The elytra lack dorsal setae except for T. otway sp. nov., which has a single dorsal seta near midlength. The parascutellar seta is present. There are 13 lateral elytral setae associated with the eighth stria, arranged in an anterior series of seven setae and a posterior series of six, and both subapical and apical elytral setae are present. As in many moriomorphines, the apical portion of interval 8 immediately laterad stria 7 is upraised as a distinct carina. Also, the metathoracic flight wings are vestigial in all Theprisa, with the metepisternum foreshortened; its maximum width subequal to greater than the lateral length.
Key to adults of Theprisa Moore  (Fig. 1A); prosternum irregularly depressed each side anterad coxal cavity, ~8 punctures associated with transverse to oblique depression; male aedeagal median lobe with broad invagination, or divot, along ventral portion of apical margin (Fig. 3A)   Pronotal base broadly punctate both on median base and laterally, laterobasal depression continuous from median base to near hind angle, with only a low tubercle laterad mesal longitudinal depression ( Fig. 2A); standardized body length smaller, 5.7-7.1 mm; male aedeagal median lobe apex broad with a straight apical face, broad surfaces of lobe apex bearing large pitted depressions (Fig. 3D)  Diagnosis (n = 5). This species is diagnosable among Theprisa spp. by the broad body and the presence of a dorsal elytral seta immediately mesad the third stria near elytral midlength (Fig. 1A). The pronotum is broad, MPW/PL = 1.38-1.46. The pronotal median base is sparsely punctate medially, with the marginal bead greatly reduced, evidenced only as a broad, slightly depressed area along the medially expanded base, in contrast to the narrowly upraised, distinct marginal bead posterad the laterobasal depressions. Whereas elytral striae 1-4 are crenulate on the disc (i.e., lined by punctures that expand the strial breadth) striae 5-7 are progressively broader and smoother, 6-7 slightly wavering along length but impunctate. In contrast T. australis, T. darlingtoni, and T. montana have striae 1-7 smooth, and T. convexa has striae 1-7 punctate near midlength. Apical abdominal ventrite of male with a single seta each side along margin, female apical ventrite with two setae each side plus a median pair of subapical setae. Standardized body length is 5.9-6.3 mm.
Description. Head narrow, ocular lobe little projected, juncture with gena very obtuse; eyes small, EyL/OLL = 0.65-0.70, not projected beyond curvature of ocular lobe, 17-20 ommatidia bisected by a horizontal line across eye; antennomeres 2 and 3 glabrous except for one dorsal seta on 2 and an apical ring of setae on 3; antennae moderately elongate, antennomere 9 maximal breadth 2× length; frontal groove deeply, medially arcuate from anterior supraorbital seta to frontoclypeal suture and continued onto lateral reaches of clypeus, area laterad groove distinctly convex; slightly concave apical labral margin 6-setose, with three smaller setae lining the lateroapical margin; mentum tooth narrowly rounded apically, sides subparallel; maxillary stipes trisetose, the three setae on the base in either a triangle with apex upward, or in an irregular horizontal line; ligula slightly convex apically, narrowed basally, trumpet shaped, its two apical setae separated by three setal diameters; paraglossae elongate, total length 2.5× distance from paraglossal base to ligular apical margin. Pronotum transverse, lateral margins straight anterad projection defined by articulatory socket of basal seta; basal margin nearly straight, the obsoletely margined median base projected posterad only slightly beyond the lateral beaded margins posterad the laterobasal depressions; median base smooth at middle, ~ 10 small punctures each side mesad deepest point of laterobasal depression; pronotal disc extended to basal marginal bead defining a tubercle that divides laterobasal depression into a median longitudinal groove and a broad lateral marginal depression inside the basal seta; median longitudinal depression deepest just anterad median base, deeply incised to very shallow, broad anterior transverse impression; anterior callosity nearly flat, front margin smooth medially, margined in outer half each side; front angles broadly, slightly protruded, subangulate; lateral marginal depression very narrow from front angle to basal 1/4 of length where it expands to meet laterobasal depression; lateral pronotal seta positioned one setal diameter inside lateral marginal depression. Prosternum medially depressed mesad anterior margins of procoxal cavities, up to 16 punctures present in a transverse band across apical 1/4 of prosternum, ~ 10 punctures each side of prosternum in depressed area anterad coxal cavity; proepisternum smooth, sutural groove with proepimeron smooth and deep. Elytra broadly hemi-ovoid, MEW/EL = 0.81-0.86, convex, sides vertically meeting lateral marginal depression; basal groove arcuate, juncture with lateral marginal depression tightly rounded, a broad, blunt tooth on margin at juncture; lateral marginal depression narrow; stria 8 deep, 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 evident, the internal plica visible ventrad deepest part of sinuation. Mesepisternum with ~ 8 shallow punctures in two vertical rows. Metepisternum wider than long, maximum width 1.8× lateral length; metepimeron fused to metepisternum laterally. Legs gracile, femora narrow, meso-and metatibiae little expanded apically, of consistent diameter throughout apical half of length; metatarsomere 1 elongate, length 0.22× tibial length, lateral sulci present on mesal and lateral faces just dorsad the ventrolateral setae. Abdominal ventrites 2-6 with 1-2 longitudinal wrinkles laterally; suture between ventrites 1 and 2 slightly curved anteriorly at midlength, ventrite 2 wrinkled within curve; suture between ventrites 2 and 3 complete laterally.
Male genitalia (n = 2). Aedeagal median lobe robust, base broadly open on right side, basal margin thickened dorsad basal opening (Fig. 3A); median lobe apex broadly rounded, slightly extended beyond ostium with a large apical divot-like concavity at tip, surface of apex densely covered with pits; internal sac bearing a stout flagellum, with apex concavely scooped (Fig. 4A), a basal articulatory sclerite associated with flagellum; right paramere elongate, parallel sided in apical half with apex narrowly rounded (Fig. 5A), bearing 12-15 short setae along ventral margin in apical half, 2-4 setae on dorsal surface near apex, though with apex glabrous; left paramere broadest near midlength, apex narrower, parallel sided with rounded tip, ventral margin bearing 2-4 setae in apical 1/3, dorsal surface with two or three setae near apex, and apex with none or one seta; antecostal apodeme of abdominal segment IX rounded distally, the apical juncture of lateral arms broad (Fig. 6A).
Female reproductive tract (n = 2). Bursa copulatrix columnar, length 1.25× maximum breadth compressed under microslide cover slip, vagina translucent, broader apical portion of bursa staining more darkly with Chlorazol black (Fig. 7A); helminthoid sclerite present, rounded apically, not extended beyond juncture with spermathecal duct; spermathecal duct stout, sinuously recurved to meet spermatheca, length subequal to length of annulated spermathecal reservoir; spermathecal gland duct very thin, length twice 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 two apical fringe setae in one dissected specimen, two setae on right gonocoxa and one on left in second dissected specimen, median surface glabrous, membranous ramus present (Fig. 8A); apical gonocoxite 2 with base extended laterally, lateral margin arcuate, apex falciform; two lateral ensiform setae and one dorsal ensiform present; two apical nematiform setae set in fossa at apical 1/4 of apical gonocoxite length.
Etymology. The species epithet is the mountain range from which this species is described, and based on current knowledge, precinctive. The epithet is to be treated as a noun in apposition.
Distribution and habitat. Theprisa otway is restricted to the Otway Ranges ( Fig. 9), with specimens collected in wet to mesic sclerophyll forest from 80-525 m elevations. The beetles are terrestrial, being discovered via raking litter, in a Berlese extraction of leaf and log litter, and via application of pyrethrin fog to logs of Eucalyptus regnans F. Mueller covered with fungus. The MCZ specimens were collected by the Darlington family in sclerophyll forest approaching rain forest in ravines, with some Nothofagus (Darlington 1960). Figures 1B, 3B Sloane stated that this species was described from specimens inhabiting "Zeehan (Simson, No. 2123); Strahan and Waratah (Carter and Lea). Eleven specimens have been examined. (Sloane 1920: 158)." We account below for the 10 paralectotypes to accompany the lectotype in the type series to ensure that this species did not include a specimen of the Tasmanian T. darlingtoni, sp. nov., described below. Paralectotypes are numbered to correspond with a gallery of images of the specimens and labels (Suppl. material 1). The first three paralectotypes are deposited in the Sloane Collection (ANIC): 1 and 2, male (left) and female (right)   Looking through registers at SAMA, we have not been able to locate this number, nor several similar missing numbers that also relate to various Tasmanian beetles from Simson.

Theprisa convexa (Sloane)
Extended diagnosis (n = 5). This species is aptly named due to the convex, domed elytra with depressed scutellar area (Fig. 1B). The elytra are narrowed basally, with the angulate humeri constricted laterally; HuW/MEW = 0.61. In keeping with the basally constricted elytra, the pronotum is more cordate than in the other species (MPW/ BPW = 1.23-1.31) and the lateral margins are sinuate anterad the pronotal basal seta articulatory socket. The pronotal base is also unique in the genus, with the marginal bead continuously marked across its breadth, and the lateral portions of the basal margin posterad the laterobasal depressions angled anteriorly relative to the extended median base. The laterobasal depression is broadly quadrate, with a moderately upraised, punctate tubercle bordered medially by a deep impression laterad the median base, and laterally by a narrow longitudinal depression inside the broadly elevated lateral margin. The body surface is glossy, with the vertex covered with indistinct transverse lines joined into an elongate transverse mesh in parts, the pronotal disc glossy with micropunctures visible across the disc, and the elytral disc and apex subiridescent due to indistinct transverse line microsculpture. Apical abdominal ventrite of male with two setae each side along margin plus two widely spaced medial subapical setae nearly in line with the outer four apical setae, female apical ventrite with two setae each side plus a transverse line of three or four medial subapical setae. Standardized body length is 6.2-7.2 mm.
Male genitalia (n = 6). Aedeagal median lobe robust, base broadly open on right side, basal margin extended as a sagittal crest dorsad basal opening (Fig. 3B); median lobe apex broadly and evenly rounded, extended only slightly beyond ostium, lateral surfaces of apex densely covered with large pits; internal sac bearing an apically narrowed lanceolate flagellum (Fig. 4B), its broadly sclerotized base extremely evident in uneverted specimens (Fig. 3B); right paramere elongate, evenly curved along length, gradually narrowed in apical half to tightly rounded tip (Fig. 5B), bearing 19-29 short setae along ventral margin in apical half, 2-9 setae on dorsal surface near apex, and with apex bearing 0-2 setae; left paramere slightly broader near midlength, apex slightly constricted before rounded apex, ventral margin usually glabrous (one individual with a single short ventral seta, a second with nine short setae), dorsal surface with 0-5 setae near apex, and apex with 0-2 setae present; antecostal apodeme of abdominal segment IX rounded distally, the apical juncture of lateral arms broad (Fig. 6B).

lectotype designations.
Extended diagnosis (n = 7). This is a large-bodied species, standardized body length 7.0-8.9 mm, with broadly based subparallel elytra, HuW/MEW = 0.67, and a cordate pronotum; the pronotal lateral margins sinuate before the hind angles (Fig. 1C). The eyes are larger than in T. australis, with ocular ratio 1.42-1.50, and 21-24 ommatidia crossed by a horizontal line bisecting the eye. The elytral striae are broad, smooth to slightly wavering along their length, and the discal elytral intervals are broadly convex. The pronotal basal marginal bead is continuous, with the margins posterad the laterobasal depressions only slightly angled forward relative to the median base. The pronotal median base bears 12-14 punctures each side isolated in a glossy surface, and the laterobasal depression is broadly, slightly tuberculate inside the broad lateral marginal depression just mesad the distinct, narrowly upraised lateral marginal bead. The body is dark, with piceous head capsule and pronotum, slightly paler rufopiceous elytra, piceous ventral surface with only the proepipleuron, elytral epipleuron, and femora dark rufous. The cuticular surface is glossy, with the vertex glossy with indistinct transverse lines in irregular wrinkles across the surface, the pronotal disc with shallow transverse mesh microsculpture, sculpticells elongate, breadth 2-4× length, and the elytra subiridescent, the surface covered with a fine, elongate transverse mesh. Apical abdominal ventrite of male with two setae each side along margin, female apical ventrite with two setae each side plus a median pair of subapical setae.
Distribution and habitat. Theprisa montana ranges in Victoria from the Dandenong Mountains, southeast to Gunyah and Tarra Valley (Fig. 9). Philip Darlington and family recorded specimens on Mt. Donna Buang at elevations ranging 450-1200 m (MCZ). One specimen was sifted from litter along a stream in Nothofagus forest at 1000 m elevation on Mt. Bullfight in the Yarra Ranges (ZMUC). Figures 2A, 3D Castelnau (1868: 198) cites the locality of this species as "Mountains of Victoria"; that locality designated type locality by Straneo (1941).

Theprisa australis (Castelnau)
Extended diagnosis (n = 5). The nearly straight, completely margined pronotal basal margin, coupled with the obtusely angulate hind angles with the pronotal lateral margins straight anterad the angles, distinguish this species from other Theprisa ( Fig. 2A). The pronotal base is broadly punctate, with 12-13 punctures each side of the median base, the punctate surface extended across the laterobasal depression to the hind angle. The elytra are moderately constricted basally, HuW/MEW = 0.64, and the elytral disc is moderately convex, the scutellum much less depressed relative to the disc than in T. convexa (Fig. 2). As in T. montana and T. darlingtoni, the elytral striae are smooth, only slightly wavering along their length. But individuals of this species are nearly always smaller, with standardized body length = 5.7-7.1 mm. The eyes are smaller than in T. montana, with ocular ratio 1.38-1.43, 15-17 ommatidia bisected by a line horizontally crossing the eye; and little convex, with the EyL/EyD ratio ranging 3.2-4.2. Body coloration is dark, with the piceous dorsal and ventral surfaces accompanied only by paler dark rufous pro-and elytral epipleura, and rufous femora. Cuticular microsculpture is well developed, with the vertex covered with an evident isodiametric mesh, and the pronotal and elytral discs covered with dense transverse lines resulting in a cyaneous iridescence. Apical abdominal ventrite of male bearing a single seta each side along margin, female apical ventrite with two setae each side plus a median group of 3-5 setae in two subapical rows.
Male genitalia (n = 5). Aedeagal median lobe robust, base broadly open on right side, basal margin sclerotized dorsad basal opening (Fig. 3D); median lobe apex broadly rounded apically, ventral margin evenly curved to meet nearly straight apical face, tip not extended beyond ostium, lateral surfaces of apex densely covered with large pits; internal sac bearing a dark field of microtrichia visible in uneverted specimens (e.g., Fig. 3D), and covering much of right side of internal sac ventrad flagellar complex (Fig. 4D); flagellum elongate, slender, with a broad base well evident in uneverted specimens (Fig. 3D); right paramere slender, evenly curved basally and narrowly extended apically to tightly rounded tip (Fig. 5D), ventral surface lined with 13-23 setae in apical 3/4 of length, 1-3 setae on dorsal surface near apex, and apex glabrous or bearing a single seta; left paramere broadest in basal half, evenly narrowed to rounded tip, ventral margin glabrous or with a single short seta near apex, dorsal surface with 0-2 setae near apex, and apex glabrous; antecostal apodeme of abdominal segment IX angled distally, lateral arms gracile, their distal juncture only slightly broader than adjoining portions of lateral arms (Fig. 6D).
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).
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 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. 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).

Discussion
Theprisa otway joins several other carabid beetle species precinctive to the Otway Ranges (Table 1). Based on biogeographic relationships among carabid beetles, area relationships of the Otways are varied, including: Tasmania alone (Stichonotus), southeastern Australia plus Tasmania (Theprisa), and eastern Victoria and New South Wales (Eutrechus, Moriomorpha, and Pseudagonica). Based on relationships among taxa in the Thayerella + Neonomius clade, the Otways represent a peripherally isolated sister-area to the remainder of temperate mainland Australia. Here again a trans-Bassian areal connection is supported, given the hypothesis that the Tasmanian monotypic genus Pterogmus Sloane is adelphotaxon to this clade (Liebherr 2019).
Zoogeographic relationships of the Otway Ranges that implicate southeastern Australia and Tasmania are also indicated for Austrocercella stoneflies, and Hemiodoecus and Hemiowoodwardia moss bugs, plus Arachnocampa glow-worms (Table 1). Geographically more restricted biogeographic relationships to taxa in South Australia and Victoria (Zephyrarchaea assassin spiders), New South Wales (Progradungula odd-clawed spiders), or eastern Victoria (Nargiotes small carrion beetles) add to the disparate examples of Otway Range biotic connections (Table 1). Finally, the Otway endemic mite, Evanssellus medusa (Lee) (Castilho et al. 2016), represents a disjunct Gondwanan pattern. Its sister species, E. foliatus (Ryke), occupies New Zealand, whereas the four species of their sister group, Heterogamasus Ryke, are distributed in southern South America (Lee 1967). Summing over these examples, the biogeographical history of the Otways has involved autochthonous speciation fragmenting ancestral ranges that include a variety of other areas of endemism, with those events arguably occurring at different times in the past. Because the relationships to these areas involve different levels of phylogenetic divergence, it is clear that the Otway area of endemism has accrued its distinct biogeographic status through various and sundry means, and thus should be considered a mosaic area of endemism that represents products of numerous different phylogenetic and biogeographic histories.
All Theprisa spp. exhibit vestigial metathoracic flight wings and dorsally convex, ovoid elytra indicating a long evolutionary history of apterous ancestors. Indeed, given the phylogenetic hypothesis proposed above (Fig. 12), Theprisa and its adelphotaxon, the clade Spherita + Sitaphe, all comprise exclusively apterous species. As such, occupation of relatively restricted geographic ranges in montane Australia (Figs 9, 10) by species with much reduced vagility is to be expected (Darlington 1936). These attributes would place Theprisa as an evolutionary terminus of the taxon pulse model (Erwin 1981). This suggests that divergence of the Tasmanian and Victorian species T. darlingtoni and T. montana respectively (Fig. 12) has not involved Pleistocene-aged trans-Bassian dispersal during low sea stands of the past 60,000 years (Orchiston 1979;Lambert and Nakada 1990), but rather an older episode of vicariance implicating presently isolated montane areas. Theprisa convexa, the other Tasmanian species stands as adelphotaxon to the other four species, thereby supporting a second, earlier episode of divergence between ancestral taxa occupying Tasmania and mainland Australia. Ad-ditionally, paraphyletic relationships among the New Zealand moriomorphine genera, Molopsida White, Tarastethus Sharp, Trichopsida Larochelle and Larivière, and Theprisa ( Fig. 12; Liebherr 2020: fig. 1) could point to either older diversification of all of these genera via fragmentation of gondwanan terranes, or more recent dispersal events across the Tasman Sea as means to establish these biogeographical disjunctions. Mutual biogeographic paraphyly is also defined by phylogenetic relationships among ancient Australian and New Zealand cave crickets (Orthoptera, Rhaphidophoridae), with estimated divergence times of these taxa arguably consistent with Gondwanan vicariance (Beasley-Hall et al. 2018;Allegrucci and Sbordoni 2019). This contribution helps build a stable classification for Moriomorphini, setting the course for additional phylogenetic research that can test whether the similar biogeographic patterns observed among moriomorphine genera are also founded on the ancient Gondwanan fragmentation of Australia, Tasmania, and New Zealand. the Darlington Collection. Catriona MacPhee is thanked for negotiating the exigencies of lockdowns, thereby providing N. Porch access to the MVM Theprisa specimens. We thank Tom Weir for his conveyed wisdom regarding the history of the Thomas G. Sloane Collection and Philip J. Darlington's, curatorial improvements of that collection during his 1957 sabbatical residency at the Australian National Insect Collection (Darlington 1960).