Hawaiian Paratachys Casey (Coleoptera, Carabidae): small beetles of sodden summits, stony streams, and stygian voids Author Liebherr, James K. https://orcid.org/0000-0001-9831-884X Department of Entomology, John H. and Anna B. Comstock Hall, 129 Garden Ave., Cornell University, Ithaca, NY 14853 - 2601, USA jkl5@cornell.edu text ZooKeys 2021 2021-06-16 1044 229 268 http://dx.doi.org/10.3897/zookeys.1044.59674 journal article http://dx.doi.org/10.3897/zookeys.1044.59674 1313-2970-1044-229 7EC23192F85D4D2AA31E2694A59CB014 0B5B533D7CBA50819E2AEC264FE6D977 Paratachys terryli sp. nov. Figures 1 , 2A , 3A , 4A , 5 Paratachys sp. "USA: Hawaii", Maddison et al. (2019 : 162). Type material. Holotype male (point-mounted, CUIC): HI: NaPali-Kona / For. Res. Kawaikoi Str. / @ Alakai Swp. Tr. 16-V- / 1991 el. 1120 m under / rocks J.K. Liebherr // HOLOTYPE ♂ / Paratachys / terryli / J. K. Liebherr 2020 (black-margined red label). Paratypes . Kauai: Halelea F. R. : Namolokama Mtn., Waioli Str., litter, sift, ohia/ferns, 22°08.00'N , 159°29.85'W , 1340 m el., 21-v-2005, lot 1 Liebherr (BPBM, 2; CUIC, 3), lot 08, Liebherr (CUIC, 1), 22°08.38'N , 159°30.18'W , 1305 m el., 23-v-2005, lot 1, Liebherr (BPBM, 2; CUIC, 3 NMNH, 2). NaPali-Kona F. R. : Alakai Swamp Tr. crossing Kawaikoi Str., 22°08.97'N , 159°36.95'W , 1130 m el., 14-v-1991, stop #91-24, Kavanaugh (CAS, 1), stop #91-26B (CAS, 1), Alakai Swamp Tr. E of Kawaikoi Str., leaf litter, sift/Berlese extraction, 22°08.85'N , 159°36.52'W , 1230 m el., 17-v-2005, lot 4, Liebherr (CUIC, 1), litter, sift/double boiler extraction, 22°08.85'N , 159°36.52'W , 1215 m el., 17-v-2005, lot 9, Liebherr (CUIC, 1; OSAC, 1); Mohihi Rdg. Tr., ohia forest litter, sift, moss, 22°06.83'N , 159°34.01'W , 1270 m el., 25-v-2005, lot 5, Liebherr (CUIC, 1); Pihea Trail, sifting leaf litter, 22°08.849'N , 159°37.889'W , 1218 m el., 26-vii-2015, Toledano & Olivieri (CTVR, 1), 29-vii-2015, Toledano & Olivieri (CTVR, 2), 1123 m, 29-vii-2015, Toledano & Olivieri (NHMUK, 2; CTVR, 3; CUIC, 2). Diagnosis. This species shares elongate elytra (EL/MEW = 1.45-1.50; Fig. 1A ) with P. perkinsi (Fig. 6B ) and P. haleakalae (Fig. 8A ), but the pronotal lateral margins are less sinuate than those of P. perkinsi , and more sinuate than those of P. haleakalae . The apical recurrent elytral groove is distinctly and narrowly impressed, with a well-defined, medially curved anterior terminus, and the elytra are truncate apically as evidenced by the nearly straight apical groove connecting the sutural and the recurrent grooves. As with P. haleakalae , elytral interneurs 1-3 are clearly impressed on the disc, but interneur 4 is traceable though discontinuous in this species, opposed to obsolete as in P. haleakalae . Standardized body length is 2.2-2.4 mm. Description. Head robust, frontal grooves shallow, convergent posterad clypeus, divergent to frontal lateral margin at frontoclypeal suture just anterad antennal articulation, broadly, slightly elevated laterally to position of anterior supraorbital seta; eyes variable, from large, macrophthalmic with 12 ommatidia crossed by horizontal diameter and 15 ommatidia crossed by vertical diameter (Fig. 1C ), to very small with dimensions of four ommatidia by five ommatidia horizontally and vertically (Fig. 1I ), with continuous variation of dimensions between those extremes (Fig. 1D-H ; see Variation section below); antennae moderately elongate, antennomere 9 ovoid, length twice diameter; labral anterior margin undulated, slightly incurved each side of midline, six-setose; penultimate maxillary palpomere broadened apically, apical palpomere a narrow spindle (Fig. 1B, E ). Prothorax transverse, MPW/PL = 1.35-1.41, base moderately constricted with lateral margins sinuate anterad right hind angles, MPW/BPW = 1.21-1.34; pronotal median base depressed, longitudinally wrinkled, unmargined; basal margin beaded each side posterad laterobasal depression which is deepest just laterad broadly triangular median base; pronotal lateral margin beaded, depression mesad bead narrow but broad enough so that a row of sculpticells can be observed lining the groove; pronotal median impression finely incised, disc flat, anterior transverse impression obsolete, not evident medially or toward slightly protruded, narrowly rounded front angles. Elytra subquadrate, lateral margins evenly convex from humeri to apex, maximum width approximately midlength; basal groove present laterad position of fifth interneur, groove convexly joined to lateral marginal depression; lateral marginal depression reflexed, of equal breadth from seta Eo2 to subapical sinuation. Pterothorax elongate, mesepisternal depression smooth posterad juncture with mesosternum, depression deepest and broadest just dorsad mesocoxal cavity; metepisternum elongate, lateral length twice maximal width; metathoracic flight wings polymorphic, 18 of 19 individuals in type series with wings reduced to a vestigial stub (Fig. 1J ), and 1 female with fully developed flight wings (Fig. 2A ), with length of alar surface 2.3 x breadth, and radial, medial, cubital and anal veins and an oblongum cell present. Abdomen with one seta each side of apical ventrite in males, two seta each side of ventrite in females. Microsculpture evident on all somites; frons covered with evident transverse mesh, sculpticells more isodiametric posteriorly on vertex; pronotum slightly iridescent due to elongate transverse sculpticells, median base and laterobasal depressions more opaque due to reticulated isodiametric microsculpture; elytra subiridescent due to a mix of elongate transverse-mesh and transverse-line microsculpture, the transverse lines denser laterally; elytral apex more opaque due to slightly raised isodiametric sculpticells between sutural interneur and apical recurrent groove; abdominal ventrites covered with elongate transverse sculpticells, the surface glossy to subiridescent. Pelage present on head, prothorax, elytra, pterothorax, abdominal ventrites and legs; pelage on head and pronotal disc, comprising microsetae separated by distances subequal to setal length (Fig. 1F, G ), as well as along ommatidial margins of the eyes (Fig. 1B ); microsetae spaced slightly farther apart on elytra, with intersetal distances up to twice microsetal length; prosternum and mesosternum medially covered with microsetae as densely distributed as on frons and pronotum; pelage on abdomen present at middle of ventrite 2 between metalegs, and progressively more broadly on ventrites 3-6 (absent dorsad arc of metaleg movement); anterior surfaces of pro-, meso-, and metathoracic legs bearing pelage of elongate microsetae, the setal bases situated more closely than microsetal lengths, trochanters and coxae similarly covered with microsetae. Coloration moderately dark; vertex and frons dark brunneous to piceous, clypeus flavobrunneous, labrum flavous; basal two antennomeres flavous, outer antennomeres progressively darker, apical segments brunneous; maxillary and labial palps flavous; pronotal disc and elytra brunneous, elytral base near scutellum paler, rufobrunneous, elytral lateral marginal depression flavobrunneous, elytral epipleuron rufoflavous, contrasted to rufobrunneous thoracic ventrites; legs flavous from trochanters outward; pro- and mesocoxae concolorous with outer leg segments, metacoxae more brunneous laterally to match the dark brunneous ventrites. Legs with basal male protarsomere alone bearing a blunt, antero-apical process. Variation. The compound eyes vary dramatically in this species, from fully macrophthalmic (Fig. 1C ), to microphthalmic (Fig. 1I ); OR ranging 1.15-1.35 for smallest- to largest-eyed individuals. That these differences reflect infraspecific variation is supported by presence of the extremes plus a variety of intermediate configurations (Fig. 1D-H ) among specimens collected microsympatrically within the identical collecting series (Fig. 1C-I ). Paratachys terryli is also polymorphic for flight-wing development, with a single macrophthalmic female specimen (Fig. 1C ) bearing fully developed flight wings (Fig. 2A ). The wings of this female are of dimensions similar to those of fully flighted individuals of Tachys oahuensis (Fig. 2B ) that were collected in ultraviolet light traps. Venation differs among the two compared species, with the wing of the female P. terryli exhibiting the oblongum cell but not the basal stem of the radius posterior ( Kukalova-Peck and Lawrence 1993 ), whereas the wing of T. oahuensis lacks the oblongum but retains the radius posterior. Both wings are folded reflexively under the elytra, with the spring-like wing margin apicad the radial cell flipping the wing open when the wing is deployed. The various brachypterous individuals exhibit a variety of eye configurations. Wing configuration is not associated with morphological variation in elytral dimensions based on comparison of four simultaneously collected females (Namolokama Mountain, 23-v-2005, lot 1) with HuW/MEW for the macropterous female = 0.57, and HuW/MEW for the three brachypterous females = 0.57-0.58. Figure 2. Right metathoracic flight wing, dorsal view A Paratachys terryli , macrophthalmic female; Kauai: Namolokama Mtn., 23-v-2005 (CUIC) B Tachys oahuensis Blackburn, female; Oahu: Honolulu, Public Health Dept. light trap, x-1965, J. W. Beardsley (BPBM). Wing vein terminology follows Kukalova-Peck and Lawrence (1993) except for the oblongum cell of Forbes (1922). From front to back of wing, veins, and cells relevant to the text include the Radial Cell (RC). Radius Anterior (RA), Radius Posterior (RP), Oblongum cell (O), Media Posterior (MP), Cubitus Anterior (CuA), Anal Anterior (AA), and Anal Posterior (AP). Male genitalia. Aedeagal median lobe porrect, parallel-sided with an evenly rounded apex (Fig. 3A ); flagellar complex elongate, with a sinuously scooped apical margin; right paramere narrow, strap-like with three apical setae; left paramere broad, elongate, with a broadly rounded apex and three apical setae. Figure 3. Paratachys spp., male aedeagus, i.e., median lobe and associated smaller right paramere and larger left paramere (right lateral view) A P. terryli ; Kauai: Namolokama Mtn., 23-v-2005 (CUIC) B P. arcanicola lectotype slide; " Tachys arcanicola Blackburn Type ♂ E.C.Z. (permanent slide, Euparal?, left paramere missing; NHMUK)." C P. haleakalae ; East Maui: Haleakala N. P., Kipahulu Vy., 30-iv-1991 (CUIC) D-F P. aaa ; Hawai'i I D Puna, Mountain View, Kazumura Cave, 21-xi-2018, lot HI00713 (CUIC) E Kau, Ocean View, Kipuka Kanohina System, 21-xi-2017, lot HI00179 (CUIC) F South Hilo, Kaumana Cave, 19-iii-2019, lot HI00942 (CUIC); disassociated left paramere below, ectal view. Female reproductive tract. Bursa copulatrix short, broad (as in Fig. 10 ), spermathecal duct narrow and extremely elongate, at least 0.57 mm long in single dissection attempt; gonocoxa bipartite, basal gonocoxite 1 narrow, elongate, with a broad, lateral sclerotized apodeme and a single apical fringe seta situated near apex of apodeme (Fig. 4A ); apical gonocoxite falcate, broadly expanded laterally at base, apex finely acuminate; two peg-like lateral ensiform setae along mid-ventral line, and one like-sized dorsal ensiform seta dorsad the apical lateral seta; two apical nematiform setae in an elongate fossa situated at approximately 2/3 length of the apical gonocoxite. Figure 4. Paratachys spp., left female gonocoxa, ventral view A P. terryli ; Kauai: Namolokama Mtn., 23-v-2005 (CUIC) B P. haleakalae ; East Maui: Haleakala N. P., Kipahulu Vy., 30-iv-1991 (CUIC) C P. aaa ; Hawai'i I.; Mountain View, Kazumura Cave, 21-xi-2018, lot HI00713 (CUIC). Abbreviations: afs, apical fringe seta; ans, apical nematiform setae; des, dorsal ensiform seta; gc1, basal gonocoxite; gc2, apical gonocoxite; les, lateral ensiform setae. Etymology. This species honors Terry L. Erwin by combining his first name and middle initial to form the genitive patronym Paratachys terryli . This construction follows that of Bembidion carlhi Erwin and Kavanaugh (1981) , a species that honors Carl H. Lindroth whose 'Carabid Beetles of Canada and Alaska' monograph (Lindroth 1969 et. seq.) revolutionized the study of North American Carabidae . Analogous to B. carlhi , this epithet recognizes Terry's immense impact on the study of Neotropical Carabidae as well as tropical biodiversity writ large. Distribution and habitat. This species is known from the Alakai Swamp west of the Wainiha River, and from Namolokama Mountain, a ridge to the east bordered by the Lumahei and Hanalei Rivers (Fig. 5 ). It has been most commonly collected in leaf and moss litter taken from low stature Ohia lehua ( Metrosideros polymorpha Gaud.; Myrtaceae ) forest. At a site at 1340 m elevation on Namolokama, one sample (21-v-2005, lot 1) sifted and subsequently hand-picked contained five Paratachys terryli in company with two Bembidion admirandum (Sharp), two B. corticarium (Sharp), and two Blackburnia kauaiensis (Sharp). A second sample from the same site (21-v-2005, lot 8) was double-boiled on a stove, resulting in discovery of one B. admirandum , eight B. corticarium , and one P. terryli . A third Namolokama sift sample from 1305 m elevation (23-v-2005, lot 1) contained seven P. terryli in company with specimens of three species of Blackburnia ; two B. posticata (Sharp), and one each B. bryophila Liebherr and B. kauaiensis (Sharp). Most other collections of P. terryli represent singletons most often collected in company with Bembidion spp., including: 16-V-1991 lot 5 from a rocky streambed also including five B. ignicola (Blackburn); 17-v-2005 lot 4 from Berlese extraction of leaf litter siftate along with two B. admirandum and three B. munroi ; and 17-v-2005 lot 9, double boiled from leaf litter siftate along with six B. admirandum and two Bl. posticata . Thus, it would appear from presently available evidence that P. terryli occurs in microhabitats most often also occupied by Bembidion beetles, either along streams, or in deep terrestrial leaf litter in montane wet ohia forest. Figure 5. Distributional records for Paratachys terryli .