Pharmacus cochleatus cochleatus ( Karny, 1935 ) cochleatus (Karny, 1935

Hegg, Danilo, Morgan-Richards, Mary & Trewick, Steven A., 2022, High alpine sorcerers: revision of the cave wētā genus Pharmacus Pictet & de Saussure (Orthoptera: Rhaphidophoridae: Macropathinae), with the description of six new species and three new subspecies, European Journal of Taxonomy 808 (1), pp. 1-58 : 26-30

Pharmacus cochleatus cochleatus ( Karny, 1935) comb. nov.

Figs 1–2 , 5–6 , 8D , 9B , 10D–E , 12D–F , 14C–D , 16A–C , 18B

Isoplectron cochleatum Karny, 1935: 383–385 , fig. 44.

Pharmacus chapmanae Richards, 1972: 158–160 , fig. 2. Syn. nov.

Pharmacus chapmanae – Chinn & Chinn 2020: 361–389.

Pharmacus brewsterensis – Chinn & Chinn 2020: fig. 4d.

Diagnosis

A mid-sized cave wētā found in the Southern Alps from Aoraki/Mt Cook south to the Dart River and east into Central Otago, at elevations greater than 1200 m. Body colour varies from chequered yellow/black to all black. Dorsal surfaces may be covered in fine tomentum, giving the insect a matt appearance, or glabrous, giving the insect a shiny appearance. The latter trait is more common at the northern end of the species’ distribution range.

At the northern end of its distribution range, P. cochleatus cochleatus overlaps with P. montanus and can easily be confused with the latter, especially at the higher elevations, where both insects are entirely black. The most reliable trait to differentiate the two species is the presence of dorsal spines on the first hind tarsus segment in P. cochleatus .

Across much of its range, P. cochleatus cochleatus overlaps with either Notoplectron brewsterense comb. nov. or with Talitropsis chopardi (Karny, 1937) east of the Main Divide of the Southern Alps. Both of the latter two species have lighter colour, shorter legs with fewer spines on the hind tibiae and no dorsal spines on the first hind tarsus segment.

Etymology

‘ Cŏchlĕăr ’ is Latin for ‘spoon’. ‘ Cochleatus ’ means ‘equipped with a spoon’ – after the shape of the male subgenital plate.

Material examined (see also Supp. file 1: Table S2 and Fig. S4 )

Holotype NEW ZEALAND • ♂, adult; locality unknown; 1876; H. Filhol leg.; MNHN EO-ENSIF4926 .

Paratype NEW ZEALAND • 1 ♂, adult; same collection data as for holotype; MNHN EO-ENSIF4928 .

Other material

NEW ZEALAND – Otago Lakes / Westland (OL/WD) • 1 ♂, holotype of Pharmacus chapmanae ; Bevan Col, Matukituki Valley ; 44.393° S, 168.689° E; 1850 m a.s.l.; Dec. 1958; M.A. Chapman leg.; OMNZ IV7927 (prev. OMNZ A70:6) • 1 ♀, allotype of Pharmacus chapmanae ; same collection data as for preceding; OMNZ IV7924 (prev. OMNZ A70:8) • 1 ♂; Bevan Col, Matukituki Valley ; 44.39424° S, 168.68760° E; 1850 m a.s.l.; 29 Jan. 2021; D. Hegg leg.; rock bluffs; night search + insect net; GenBank: OM293736 ; MPN CW5179 • 1 ♂, 1 ♀; same collection data as for preceding; MPN CW5178 , CW5180 . – Westland (WD) • 1 ♀; Topheavy, Mt Brewster, Haast Pass ; 44.06548° S, 169.42843° E; 2000 m a.s.l.; 18 Feb. 2017; D. Hegg leg.; on rocky ridge above glacier; night search; GenBank: OM293694 ; MPN CW3322 • 2 ♂♂, 5 ♀♀; same collection data as for preceding; MPN CW3277 , CW3279 to CW3282 , CW3324 , CW3325 . – Otago Lakes (OL) • 1 ♀; Luna Basin, Major Peak, Wakatipu ; 44.92490° S, 168.47476° E; 1470 m a.s.l.; 9 Apr. 2016; D. Hegg leg.; on large rocks in alpine basin; night search; GenBank: OM293686 ; MPN CW2985 • 3 ♂♂, 1 ♀; same collection data as for preceding; MPN CW2979 , CW2981 , CW2999 , CW3000 • 1 nymph; Coronet Peak, Queenstown ; 44.91774° S, 168.73423° E; 1540 m a.s.l.; 18 Jan. 2019; D. Hegg leg.; on rock formations; night search + insect net; GenBank: OM293709 ; MPN CW4288 • 5 ♂♂, 3 ♀♀; same collection data as for preceding; MPN CW4414 to CW4418 , CW4432 to CW4434 • 1 ♀; Lochnagar, Richardson Mountains ; 44.59627° S, 168.56400° E; 2080 m a.s.l.; 21 Sep. 2019; D. Hegg leg.; on rocks on side of snow couloir; casual find while climbing; GenBank: OM293726 ; MPN CW4590 • 1 ♂; Sentinel Peak, Lake Hāwea ; 44.41029° S, 169.24003° E; 1580 m a.s.l.; 5 Dec. 2020; D. Hegg leg.; on rocky ridge; night search + insect net; GenBank: OM293731 ; MPN CW5134 • 2 ♂♂, 2 nymphs; same collection data as for preceding; MPN CW5132 , CW5133 , CW5135 , CW5136 • 2 ♂♂; Earnslaw Burn ; 44.67482° S, 168.39371° E; 1280 m a.s.l.; 13 Nov. 2020; D. Hegg leg.; on rocky ridge and cliffs; night search + insect net; GenBank: OM293733 , OM293734 ; MPN CW5149 , CW5150 • 1 ♂, 1 ♀, 2 nymphs; same collection data as for preceding; MPN CW5151 to CW5154 • 1 ♀; Roys Peak, Wānaka ; 44.69477° S, 169.04644° E; 1500 m a.s.l.; 19 Nov. 2020; D. Hegg leg.; rock bluffs; night search + insect net; GenBank: OM293732 ; MPN CW5137 • 5 ♂♂, 2 ♀♀, 2 nymphs; same collection data as for preceding; MPN CW5138 to CW5144 , CW5147 , CW5148 • 1 ♀, 1 nymph; below Bevan Col, Matukituki Valley ; 44.39546° S, 168.68503° E; 1750 m a.s.l.; 29 Jan. 2021; D. Hegg leg.; rock slabs; casual find while climbing; MPN CW5181 , CW5182 • 1 nymph; Turret Head, Earnslaw Burn ; 44.65757° S, 168.37977° E; 2300 m a.s.l.; 2 Feb. 2021; D. Hegg leg.; rocky ridge; casual find while climbing; GenBank: OM293735 ; MPN CW5176 • 1 nymph; same collection data as for preceding; MPN CW5175 • 3 ♂♂, 1 ♀, 1 nymph; Turret Head, Earnslaw Burn ; 44.65681° S, 168.38646° E; 2100 m a.s.l.; 2 Feb. 2020; D. Hegg leg.; rock bluffs; night search + insect net; MPN CW5244 to CW5247 , CW5177 • 1 ♂, 2 ♀♀, 5 nymphs; Albertburn Saddle, East Matukituki ; 44.39752° S, 168.87310° E; 1600 m a.s.l.; 5 Feb. 2021; D. Hegg leg.; rock bluffs; night search + insect net; MPN CW5170 to CW5174 , CW5238 to CW5240 • 2 ♂♂, 1 ♀; East Matukituki / Albertburn divide ; 44.36994° S, 168.8887° E; 1830 m a.s.l.; 6 Feb. 2021; D. Hegg leg.; rock bluffs; night search + insect net; MPN CW5241 to CW5243 • 2 ♀♀; French Ridge, West Matukituki ; 44.42520° S, 168.69874° E; 1800 m a.s.l.; 20 Feb. 2021; D. Hegg leg.; rocky ridge; night search + insect net; MPN CW5207 , CW5208 . – Central Otago (CO) • 1 ♂; Mt Pisa, Pisa Range ; 44.88278° S, 169.19500° E; 1650 m a.s.l.; 27 Feb. 2018; D. Hegg leg.; on tor; night search + insect net; GenBank: OM293705 ; MPN CW3831 • 4 ♂♂, 5 ♀♀, 2 nymphs; same collection data as for preceding; MPN CW3795 to CW3797 , CW3828 to CW3830 , CW3847 to CW3851 . – Mackenzie (MK) • 1 ♀; Dasler Pinnacles, Naumann Range ; 43.95237° S, 169.86241° E; 1800 m a.s.l.; 14 Jan. 2017; D. Hegg leg.; on rock bluffs; night search; GenBank: OM293691 ; MPN CW3284 • 3 ♂♂; same collection data as for preceding; MPN CW3283 , CW3285 , CW3296 • 1 ♂; Mt Edgar Thomson, Ben Ōhau Range ; 43.78252° S, 170.05561° E; 2350 m a.s.l.; 28 Jan. 2017; D. Hegg leg.; on mixed rock and snow ridge; casual find while climbing; GenBank: OM293690 ; MPN CW3278 .

Description

MEASUREMENTS. See Table 1.

HEAD. As per generic description. Eyes dark. Specimens living above the permanent snowline are entirely black. This includes all head parts that are usually pale otherwise, e.g., maxillary palps and antennae.

THORAX. As per generic description.

LEGS. Notably shorter than in P. montanus . The hind tibiae are approx. 10% shorter than the body in both males and females. Fore femora always unarmed at the apex. Mid femora armed with one retrolateral spine at the apex; prolateral apical spine absent. The first hind tarsus segment is armed with a variable

number of linear spines above (up to nine); the second hind tarsus segment is only rarely armed with dorsal linear spines. The first tarsal segment on all six legs is pale at times, but not always; the insect can be entirely black ( Fig. 14D ).

ABDOMEN. Tergites tomentose or glabrous; the latter trait appears to be more common in the northern half of the range. A thin, pale median line along the whole length of the insect may or may not be present. The colour of the tergites is mostly chequered yellow/black or light grey-brown/black at the lower elevations ( Fig. 14C ), transitioning to mostly black ( Fig. 8D ) or all black ( Fig. 14D ) at the higher elevations, especially above the permanent snow-line.

MALE TERMINALIA. Subgenital plate an isosceles triangle with very rounded corners and a spoon on the vertex; keeled and mostly glabrous, with some sparse hair near the edges; shallow when seen from the side. In a dorsal view, only the spoon is visible, protruding beyond the paraprocts; these are covered in short, dense bristles ( Fig. 10D–F ).

FEMALE TERMINALIA. Subgenital plate bilobed, the two rounded lobes short, broad, and asymmetrical, with a wide gap in between; very hairy ( Fig. 12D ). Ovipositor on average 70% of body length, only very gently curving upwards; lower valve with 5 to 9 strong teeth below at the apex ( Fig. 12E–F ).

Figures

Fig. 1. A. Genetic relationships of cave wētā support monophyly of the genus Pharmacus Pictet & de Saussure, 1893. Gene tree from mtDNA sequences (~1500 bp of COI) using Maximum Likelihood with 1000 bootstraps and Macropathus filifer Walker, 1869 to root the tree. B. Map of South Island of New Zealand, showing locations of Pharmacus specimens used in phylogenetic analyses (Figs 1A, 2). Colours correspond to different species. Two letter codes indicate the New Zealand entomological regions (Crosby et al. 1998). These codes are reported in the Material examined section for each species.

Fig. 2. Gene tree for the eight morphologically identified species of Pharmacus Pictet & de Saussure, 1893 using Maximum Likelihood analysis of ~850 bp of mtDNA (COI) from 60 specimens ofPharmacus.

Fig. 5. Pharmacus cochleatus (Karny, 1935) comb. nov. A–D. Adult ♂. Dorsal view of terminalia. A. Holotype, from unknown location (MNHN EO-ENSIF4926). B. Paratype, from unknown location (MNHN EO-ENSIF4928). C. Original drawing by Aola Richards (1972): Pharmacus chapmanae Richards, 1972 holotype. Bevan Col, Matukituki Valley (OMNZ IV7927; prev. OMNZ A70:6). D. Mt Edgar Thomson, Ben Ohau Range (MPN CW3278). E–F. Adult ♀. Ventral view of terminalia (subgenital plate). E. Original drawing by Aola Richards (1972): Pharmacus chapmanae Richards, 1972 allotype. Bevan Col, Matukituki Valley (OMNZ IV7924; prev. OMNZ A70:8). F. Mt Brewster, Haast Pass (MPN CW3325). Scale bar = 2 mm. Images A and B courtesy of Marion Depraetere, MNHN, reproduced under CC BY-NC-ND 4.0 licence.

Fig. 6. Known distribution of cave wētā in the genus Pharmacus Pictet & de Saussure, 1893 in the south of the South Island, New Zealand.

Fig. 8. Dorsal views of adult ♂ cave wētā in the genus Pharmacus Pictet & de Saussure, 1893. Notice loss of natural pigmentation of pale body parts due to preservation in ethanol. A–B. Pharmacus montanus Pictet & de Saussure, 1893 A. Mt Annette, Sealy Range, Mt Cook (MPN CW3302). B. Lake Anna, Mt Franklin, Arthur’s Pass (MPN CW3989). C. Pharmacus senex sp. nov., Old Woman Range, Central Otago (MPN CW4387). D. Pharmacus cochleatus cochleatus (Karny, 1935) comb. nov., Topheavy, Mt Brewster, Haast Pass (MPN CW3324). E. Pharmacus cochleatus rawhiti subsp. nov., Mt Tūwhakarōria, Hector Mountains (NMNZ AI.052290). F. Pharmacus cristatus sp. nov., Skippers Range, South Westland (MPN CW4562). G. Pharmacus vallestris sp. nov., Matukituki River West Branch (MPN CW3700). H. Pharmacus notabilis sp. nov., Remarkables ski-field access road (NMNZ AI.052296). I. Pharmacus concinnus sp. nov., Eyre Peak, Eyre Mountains (MPN CW4482). J. Pharmacus perfidus sp. nov., Spence Peak, Takitimu Mountains (MPN CW4494). Scale bar = 10 mm.

Fig. 9. Left hind tibia of adult ♂ cave wētā in the genus Pharmacus Pictet & de Saussure, 1893, dorsal view. A. Pharmacus montanus Pictet & de Saussure, 1893, Lake Anna, Mt Franklin, Arthur’s Pass (MPN CW3989). B. Pharmacus cochleatus cochleatus (Karny, 1935) comb. nov., Gertrude Saddle, Darran Mountains (MPN CW3413). C. Pharmacus cochleatus rawhiti subsp. nov., Mt Tūwhakarōria, Hector Mountains (MPN CW4431). D. Pharmacus cristatus sp. nov., Skippers Range, South Westland (NMNZ AI.052292). E. Pharmacus notabilis sp. nov., Remarkables ski-field access road (NMNZ AI.052296). F. Pharmacus senex sp. nov., Obelisk, Old Man Range (NMNZ AI.052294). G. Pharmacus concinnus sp. nov., Eyre Peak, Eyre Mountains (MPN CW4480). H. Pharmacus perfidus sp. nov., Spence Peak, Takitimu Mountains (NMNZ AI.052300). I. Pharmacus vallestris sp. nov., Matukituki River West Branch (MPN CW3700). Scale bar = 5 mm.

Fig. 10. Adult male terminalia of cave wētā in the genus Pharmacus Pictet & de Saussure, 1893. Left column: dorsal view; central column: ventral view (subgenital plate); right column: lateral view. A–C. Pharmacus montanus Pictet & de Saussure, 1893, Mueller Hut Track, Sealy Range, Mt Cook (MPN CW3532). D–F. Pharmacus cochleatus (Karny, 1935) comb. nov. D–E. Humboldt Mountains (MPN CW3698). F. Blue Lake Creek, Garvie Mountains (MPN CW4462). G–I. Pharmacus cristatus sp. nov., Skippers Range High Point, South Westland (NMNZ AI.052292). J–L. Pharmacus notabilis sp. nov., Remarkables Ski-field access road (NMNZ AI.052296). Scale bars = 1 mm.

Fig. 12. Adult female terminalia of cave wētā in the genus Pharmacus Pictet & de Saussure, 1893. Left column: subgenital plate; central and right columns: ovipositor. A–C. Pharmacus montanus Pictet & de Saussure, 1893. A. Mt Annette, Sealy Range, Mt Cook (MPN CW3303). B–C. Mt Wakefield, Mount Cook Range (MPN CW3362). D–F. Pharmacus cochleatus (Karny, 1935) comb. nov., Lochnagar, Richardson Mountains (MPN CW4590). G–I. Pharmacus cristatus sp. nov., Skippers Range High Point, South Westland (NMNZ AI.052293). J–L. Pharmacus notabilis sp. nov., Two Mile Hut, Hector Mountains (NMNZ AI.052297). Scale bars = 1 mm.

Fig. 14. Live Pharmacus Pictet & de Saussure, 1893 in their natural environments. A–B. Pharmacus montanus Pictet& de Saussure, 1893.A.Adult ♂, with dark coloration typicalof high elevationspecimens. Mt Annette, Sealy Range, Mt Cook, 2200 m a.s.l. (MPN CW3302). B. Adult ♂, with coloration typical of low elevation specimens. Mueller Hut Track, Sealy Range, Mt Cook, 1500 m a.s.l. C–D. Pharmacus cochleatus cochleatus (Karny, 1935) comb. nov. C. Adult ♀, with yellow / black chequered coloration that is most typical for the species, parasitized by red mites (Order Trombidiformes). Luna Basin, Major Peak, Wakatipu, 1470 m a.s.l. D. Adult ♂, with dark coloration typical of high elevation specimens. Mt Edgar Thomson, Ben Ōhau Range, 2350 m a.s.l. (MPN CW3278). E. Pharmacus cochleatus rawhiti subsp. nov. Adult ♂, Blue Lake Creek, Garvie Mountains, 1280 m a.s.l. F. Pharmacus cristatus sp. nov. Adult ♀, Skippers Range High Point, South Westland, 1520 m a.s.l.

Fig. 16. Habitat of Pharmacus Pictet & de Saussure, 1893.A. Sealy Range, Mount Cook.a–b. Collection localities of Pharmacus montanus Pictet & de Saussure, 1893. a. Mueller Hut Track, 1500 m a.s.l. (MPN CW3299, CW3300, CW3532–3535). b. Mount Annette, 2200 m a.s.l. (MPN CW3302, CW3303, CW3323). c. Mount Edgar Thomson, Ben Ohau Range, 2350 m a.s.l., collection locality of P. cochleatus cochleatus (Karny, 1935) comb. nov. (MPN CW3278). B. Lochnagar, Richardson Mountains. d. Rock bluffs at 2080 m a.s.l., collection locality of P. cochleatus cochleatus (Karny, 1935) comb. nov. (MPN CW4590). C. Matukituki River West Branch. e–f. Collection localities of P. vallestris sp. nov. e. Valley floor, dry creek bed at 1000 m a.s.l. (MPN CW3430, CW3700). f. French Ridge, 1650 m a.s.l. (MPN CW5145, CW5146, CW5213–5215). g–h. Collection localities of P. cochleatus cochleatus (Karny, 1935) comb. nov. g. French Ridge, 1800 m a.s.l. (MPN CW5207, CW5208). h. Bevan Col, 1850 m a.s.l. (OMNZ IV7927 and MPN CW5178 to 5180). D. The Remarkables, Queenstown. i–j. Collection localities of Pharmacus notabilis sp. nov. i. Remarkables Ski-field Access Road, 1220 m a.s.l. (NMNZ AI.052296). j. The Remarkables North ridge, 1600 m a.s.l. (MPN CW2588 – 2594). k. Lake Alta, 1820 m a.s.l., collection locality of Pharmacus cochleatus rawhiti subsp. nov. (MPN CW5219 – 5221).

Fig. 18. Map of South Island, New Zealand, showing the known distribution of cave wētā in the genus Pharmacus Pictet & de Saussure, 1893. A–B. Solid shapes indicate material collected as part of this study; empty circles indicate additional locations of material examined by Richards (1972). B. Shapes represent different subspecies: dark blue circle = Pharmacus cochleatus cochleatus; light blue circle = Pharmacus cochleatus nauclerus subsp. nov.; diamond = Pharmacus cochleatus fiordensis subsp. nov.; triangle = Pharmacus cochleatus rawhiti subsp. nov.

Fig. 4. Side by side comparison of Notoplectron brewsterense (Richards, 1972) comb. nov. and of Pharmacus montanus Pictet & de Saussure, 1893. A–E, H, J. Notoplectron brewsterense (Richards, 1972) comb. nov., Mt Brewster, Haast Pass. A–C. Dorsal, ventral and lateral view of terminalia. Adult ♂ (MPN CW3307). D–E. Subgenital plate. Adult ♀. D. MPN CW3391. E. Original drawing by Aola Richards (1972): holotype (CMNZ 000267). H, J. Dorsal view of hind tibia with enlarged detail.Adult ♀ (MPN CW3391). F–G, I, K. Pharmacus montanus Pictet & de Saussure, 1893. F–G. Subgenital plate. Adult ♀. F. Mt Annette, Sealy Range (MPN CW3303). G. Original drawing by Aola Richards (1972), from unspecified material. I, K. Dorsal view of hind tibia with enlarged detail. Adult ♂. Lake Anna, Mt Franklin, Arthur’s Pass (MPN CW3989). Scale bars = 1 mm.