Pharmacus montanus Pictet & de Saussure, 1893

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 : 20-25

Pharmacus montanus Pictet & de Saussure, 1893

Figs 1–3 , 4F–G, I, K , 7 , 8A–B , 9A , 10A–C , 12A–C , 14A–B , 16A , 18A

Pharmacus montanus Pictet & de Saussure, 1893: 302–303 , pl. I fig. 5–5c.

Pharmacus dumbletoni Richards, 1972: 161–162 , fig. 4. Syn. nov.

Pharmacus montanus – Hutton 1896: 239. — Richards 1972: 156–158, fig. 1.

Diagnosis

A mid-sized cave wētā found in the Southern Alps from the Ben Ōhau Range northwards, at elevations greater than 1200 m. Body colour varies from marbled brown to black, generally with a thin but well visible median dorsal line along the whole length of the animal, and pale first tarsal segments. The pronotum and tergites are always covered in fine tomentum, giving the animal a matt appearance.

At the southern end of its distribution range, Pharmacus montanus overlaps with P. cochleatus comb. nov. and could be easily confused with the latter, especially at the higher elevations, where both insects are black. P. cochleatus is often glabrous and shiny in appearance and has first segment of the hind tarsi armed with dorsal linear spines, which are absent in P. montanus .

On the higher mountain tops east of the Main Divide of the Southern Alps, P. montanus shares the habitat with Petrotettix serratus Richards, 1972 . The latter species is often dark with pale tarsi like P. montanus but has dorsal linear spines on the first two segments of the hind tarsi, and a serrated upper valve of the ovipositor.

In the alpine regions around Mt Cook, P. montanus forms mixed populations with Notoplectron brewsterense comb. nov. The latter is yellow-orange in colour, has a stocky appearance due to its shorter legs, and has fewer, larger dorsal linear spines on the hind tibiae. See also Fig. 4 for a comparison between the two species.

At the northern end of its distribution range in Kahurangi National Park, P. montanus is uniform brown and shares the habitat with Macropathus filifer , which is also brown. The latter species however has much longer legs, which give it a very slender appearance.

Etymology

‘ Pharmăcus montānus ’ is Latin for ‘mountain sorcerer’ – an inspired name for Pictet’s and de Saussure’s (1893) “entirely black insect”.

In common English, P. montanus is also known as ‘Mount Cook flea’. The name is explained by the insect’s “reprehensible habit of leaping in showers out of a crevice in the rock upon the unsuspecting climber” ( Dumbleton 1935), and is still in common use today.

Material examined (see also Supp. file 1: Table S1 and Figs S2–3 )

Holotype NEW ZEALAND • ♂, adult; Mackenzie (MK), Mt Cook region; 43.75° S, 170.06° E; 2100 m a.s.l.; date unknown; G. Mannering leg.; MHNG ARTO-24289 .

Other material

NEW ZEALAND – Mackenzie (MK) • 1 ♀; Mt Annette, Sealy Range ; 43.75129° S, 170.06210° E; 2200 m a.s.l.; 25 Feb. 2017; D. Hegg leg.; summit rocks, above glacier; night search; GenBank: OM293693 ; MPN CW3303 • 1 ♂, 1 ♀; same collection data as for preceding; MPN CW3302 , CW3323 • 1 ♂; Mueller Hut Track, Sealy Range ; 43.71504° S, 170.06982° E; 1500 m a.s.l.; 24 Feb. 2017; D. Hegg leg.; under large boulder; night search + insect net; GenBank: OM293692 ; MPN CW3300 • 3 ♂♂, 2 nymphs; same collection data as for preceding; MPN CW3299 , CW3532 to CW3535 • 1 nymph; Mt Wakefield, Mount Cook Range ; 43.70707° S, 170.12170° E; 1660 m a.s.l.; 18 Mar. 2017; D. Hegg leg.; on rock bluffs; night search + insect net; GenBank: OM293695 ; MPN CW3330 • 4 ♀♀, 1 nymph; same collection data as for preceding; MPN CW3331 , CW3406 , CW3660 to CW3662 • 1 ♀, 1 nymph; Mt Wakefield, Mt Cook Range ; 43.71989° S, 170.12771° E; 1750 m a.s.l.; 13 May 2017; D. Hegg leg.; on rocky ridge; night search + insect net; MPN CW3516 , CW3699 • 1 nymph; Mt Dark, Ben Ōhau Range ; 43.87117° S, 170.04536° E; 1800 m a.s.l.; 17 May 2020; D. Hegg leg.; under boulder on scree slope; casual find while climbing; GenBank: OM293728 ; MPN CW4876 • 1 nymph; same collection data as for preceding; MPN CW4875 . – North Canterbury/Westland (NC/WD) • 1 nymph; Lake Anna, Mt Franklin, Arthur’s Pass ; 42.87567° S, 171.65048° E; 1720 m a.s.l.; 23 Dec. 2017; D. Hegg leg.; on rock bluffs; night search; GenBank: OM293702 ; MPN CW3650 • 1 ♂, 1 ♀; same collection data as for preceding; MPN CW3988 , CW3989 . – Buller ( BR) • 1 ♂, holotype of P. dumbletoni ; Mount Mahanga, Spenser Mountains ; 42.102° S, 172.635° E; 2100 m a.s.l.; Oct. 1950; L.J. Dumbleton leg.; NZAC 03015679 • 1 ♂; Gloriana Peak, Spenser Mountains ; 42.27437° S, 172.48992° E; 1750 m a.s.l.; 5 Feb. 2017; D. Hegg leg.; on rock bluffs; night search + insect net; GenBank: OM293689 ; MPN CW3262 • 1 ♂, 1 ♀, 2 nymphs; same collection data as for preceding; MPN CW3263 to 3266 • 4 ♂♂, 2 ♀♀; same collection data as for preceding; 25 Apr. 2017; MPN CW3656 to CW3659 , CW3665 , CW3666 • 1 ♀; Between Sunset Saddle and Mt Hopeless, Travers Range ; 41.91926° S, 172.73519° E; 1900 m a.s.l.; 31 Dec. 2020; D. Hegg leg.; in rocky chasm; visual search in daylight; MPN CW5183 . – Nelson (NN) • 1 ♀; Poverty Basin, Mt Owen ; 41.54712° S, 172.52864° E; 1640 m a.s.l.; 8 Jan. 2019; D. Hegg leg.; on limestone bluffs; night search + insect net; GenBank: OM293708 ; MPN CW4275 • 4 ♂♂, 1 ♀, 4 nymphs; same collection data as for preceding; MPN CW4272 , CW4273 , CW4446 to CW4450 , CW4456 , CW4457 .

Description

MEASUREMENTS. See Table 1.

HEAD. As per generic description. 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; always covered in fine hair, which gives the animal a matt look.

LEGS. Longer than in most Pharmacus species. The hind tibiae are 5% longer than the body in females, 20% longer than the body in males. Fore femora always unarmed at the apex. Mid femora armed with one retrolateral spine at the apex; prolateral apical spine absent. Hind femora occasionally armed with up to two linear spines below, on either or both inner and outer edge. Hind tibiae armed with two superior subapical spines (one prolateral and one retrolateral), two superior apical spines (one prolateral and one retrolateral), two inferior apical spines (one prolateral and one retrolateral) and two inferior subapical spines (one prolateral and one retrolateral), as in all other Pharmacus species. However, in some specimens in the Mt Cook region, a third inferior subapical spine may be present on the inner edge of the tibia (see Fig. 3F–G ). First and second tarsal segments without any linear spines above – a trait that differentiates P. montanus from all other Pharmacus species. The first tarsal segment is generally pale on all six legs, even in specimens that are entirely black otherwise – the contrast is in fact especially noticeable in the darker specimens and gives the insect a characteristic look ( Fig. 14A ).

ABDOMEN. Tergites always covered in fine hair, which gives the animal a matt look. A thin, pale median line along the whole length of the insect is generally visible, even in the darker specimens. The colour of the tergites is most commonly brown at the lower elevations ( Fig. 14B ), transitioning to black at the higher elevations, especially above the permanent snow-line ( Figs 8A , 14A ). Specimens in the Arthur’s Pass region are chequered, unlike other P. montanus but like other Pharmacus species further south ( Fig. 8B ).

MALE TERMINALIA. Subgenital plate triangular when seen from below; deeper than in all other Pharmacus species when seen from the side ( Fig. 10A–C ). In a dorsal view, a thin plate or membrane covered in tomentum is attached to the vertex of the triangle. The plate varies in shape from oval ( Figs 3D , 8B ) to rectangular with rounded corners ( Figs 3E , 10A ).

FEMALE TERMINALIA. Subgenital plate strongly bilobed, the two rounded lobes long and narrow, separated by a deep V-shaped depression; hairy ( Figs 4F–G , 12A ). Ovipositor on average 70% of body length, relatively straight, tapering gently at first then more strongly near the apex; lower valve with 5 to 8 strong teeth below at the apex ( Fig. 12B–C ).

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. 3. Pharmacus montanus Pictet & de Saussure, 1893. Adult ♂. A–B. Lateral view of terminalia. A. Original drawing by Pictet & de Saussure (1893): Holotype, Mt Cook (MHNG ARTO-24289). B. Mt Annette, Sealy Range, Mt Cook (MPN CW3302). C–E. Dorsal view of terminalia. C. Original drawing by Pictet & de Saussure (1893): holotype, Mt Cook (MHNG ARTO-24289). D. Original drawing by Richards (1972), from unspecified material. E. Mt Annette, Sealy Range, Mt Cook (MPN CW3302). F–G. Detail of ventral spines at apex of left hind tibia. F. Mt Annette, Sealy Range, Mt Cook (MPN CW3302), with two sub-apical spines on inner edge and one on the outer edge as per the species’ original description. G. Lake Anna, Mt Franklin, Arthur’s Pass (MPN CW3989), with one sub-apical spine on both inner and outer edges, a configuration typical of all Pharmacus Pictet & de Saussure, 1893. Scale bars = 1 mm.

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.

Fig. 7. Head of Pharmacus montanus Pictet & de Saussure, 1893, Lake Anna, Mt Franklin, Arthur’s Pass. A–B. Adult ♂ (MPN CW3989) C–D. Adult ♀ (MPN CW3988). Scale bar = 2 mm.

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.