Kuschelysius , a New Alpine Genus of Eugnomine Weevil ( Coleoptera : Curculionidae : Curculioninae ) from New Zealand

Kuschelysius new genus is described for four species, K. hollowayae new species, K. durus new species, K. verbalis new species and K. nitens new species, which are found in alpine regions along the length of the South Island of New Zealand. The genus most closely resembles members of the genus Eugnomus but is distinguished from them by the presence of a small pair of post-ocular tubercles and by having appressed scales on the dorsal surfaces. Some members of Kuschelysius appear to be flight-capable with well-developed hindwings, while others have reduced hindwings and are presumably flightless. Many specimens have been collected from the flowers of Dracophyllum traversii, Celmisia and other alpine plants, and the guts of examined specimens contained pollen. We hypothesise that the species of Kuschelysius are pollinators of the New Zealand alpine flora.


Introduction
The Eugnomini Lacordaire, 1863 are a tribe of curculionine weevils with a primarily Southern Hemisphere distribution.The tribe is present in Australia, New Zealand, New Caledonia, South America extending northward to Central America, with undescribed members also known from Indochina [1].The Eugnomini reach their greatest diversity in New Zealand, with approximately 100 species in 19 endemic genera, one genus (Rhopalomerus Blanchard) shared with Australia and South America and another (Pactola Pascoe) shared with New Caledonia and Fiji [2].They form a distinctive element of the New Zealand weevil fauna, where their diurnal activity, often bizarre morphologies masquerading as faeces while feigning death, and flower-frequenting behaviour [3,4] attract the attention of natural historians.
The exact composition of the Eugnomini remains unclear.A suite of characters has been used to characterise the tribe.These characters include an elongate head with the eyes separated from the head constriction by a distance greater than their own length, scrobes that run below the lower surface of the rostrum, front coxae conical, and a large, obvious tooth on the hind tibia [3]: some of these characters are not universally shared and there have not yet been any phylogenetic studies to identify synapomorphies that unequivocally indicate the monophyletic nature of the tribe [1].Substantial advances have been made in recent years on the eugnomine weevil fauna of New Caledonia and the Pacific [2,[5][6][7][8][9], but the remainder of the Eugnomini of the world remain understudied.
Despite the lack of regional or global revisions of the tribe, the lowland eugnomine weevils of New Zealand are reasonably well known and have been named and inventoried through Broun's pioneering taxonomic research [10][11][12][13][14], illustrated accounts of several species by Hudson [15,16] and a key to genera by Marshall [17].However, it is becoming apparent that the diversity of weevils in alpine areas has yet to be fully appreciated, with many species belonging to several genera being found exclusively above the treeline [18].
This contribution describes a new genus of eugnomine weevil containing four distinctive, large, alpine species, which we dedicate to our late friend, mentor and colleague Guillermo (Willy) Kuschel.We also name the type species after our friend and colleague Beverly Holloway, Willy's wife, a leading coleopterist in her own right.Although Willy's published contributions on the Eugnomini were only ever part of broader works [19][20][21][22], he was fascinated by the group and worked on the fauna of New Zealand during his first lengthy visit to the country and soon after his permanent settlement in New Zealand.Regrettably, the results from this research were not published in his lifetime.

Materials and Methods
Morphological features are described using standard terminology [23][24][25], with the following additions.Body length was measured in lateral view, from the anterior margin of the eyes to the elytral declivity.Body height was measured in lateral view as a straight line from the hind coxae to the dorsal surface of the elytra.Rostrum width was measured across the antennal insertions.The curve formed by the connection of the head and rostrum, as observed in lateral view, is termed the "ventral curvature of the head".The degree of curvature can be described as "gently curved" (i.e., having a large circle of curvature) or "tightly curved" (i.e., having a small circle of curvature).Wing venation terminology follows Kukalová-Peck and Lawrence [26], as applied to Curculionoidea by Oberprieler et al. [23].This relates to the terminology of Zherikhin and Gratshev [27] in the following way: C = C; Sc = Sc; RA = R; RC = w; rf = rf; RP = Rr; RP 1 = pst; RP 2 = h & mst; rs = rs; MP 1+2 = Cu; msc = msc; ms = af; MP 3 = 1A 1 ; MP 4 = 1A 2 ; CuA = 2A; CuA 1 = a 1 -a 2 AA = 3A; ac = ac; AP = 4A; J = J.
Descriptions of colour follow the terminology provided by the National Bureau of Standards [28], which gives 267 centroid colours with natural-language descriptions.Digital representations of these colours have been provided by Jaffer [29].
Genitalia were photographed in KY Jelly (Johnson & Johnson Pacific, Broadway, NSW, Australia) before being stored in glycerol in a vial pinned below the specimen.Illustrations were prepared from photographs, using Inkscape (v.0.91, [30]).Habitus photographs were taken using Nikon DS-Ri1 (Melville, NY, USA) fitted with a digital camera and a mechanical z-stepper.Nikon NIS Elements v. 4.10 was used to prepare the image stack and to produced the final montaged image.
Specimens were examined and deposited in the following collections: Label data from holotypes are transcribed using the following conventions.Data from individual labels are enclosed using quotes ('. . .'), lines are indicated with a solidus (/) and metadata are given in square brackets ([. . .]). Two-letter regional codes (NN, BR, FD, etc., Figure 1) follow those proposed by Crosby et al. [31].

Kuschelysius Brown and Leschen New Genus
Type species: Kuschelysius hollowayae Brown and Leschen new species, by present designation.

Diagnosis
Large-bodied (body length > 5.0 mm, height > 1.7 mm, width > 2.2 mm); dorsal surfaces covered with appressed scales; antennal club with segment 3 shorter than segments 1 and 2 combined; length of the rostrum longer than the head; head not constricted behind eyes, tubercles present behind eyes; distinct spines absent from pronotum and elytra; elytra with a humeral callus; all femora with a single ventral tooth; all tibiae sinuous; tarsal claws simple, lacking a tooth.
The presence of a pair of small tubercles present behind the eyes will distinguish Kuschelysius from most other genera of New Zealand eugnomine weevils.Kuschelysius is most similar to species of Eugnomus Schönherr: the presence of appressed scales which conceal the integument, and sinuous proand mesotibiae distinguish them from Eugnomus, which have dorsal vestiture of fine hairs that reveal the integument, head evenly convex behind the eyes and straight pro-and mesotibiae.Tysius Pascoe also has tubercles behind the eyes; however, this genus can be distinguished from Kuschelysius by its much smaller size (body length < 3 mm), eyes placed further onto the rostrum, round scutellar shield and by the elytra having facia on interstria 3 and a low tubercle on interstria 5 on the elytral declivity.

Description
Rostrum.Mandibles stout, not exodont.Maxillae with long and flexible palps.Antennae inserted laterally, at distal 1/4 of rostrum.Scrobes oblique, running along ventral surface for 2/3 length, terminating just short of eyes.Head.Eyes hemispherical, prominent, positioned anterior of point of maximum ventral head curvature.Head not constricted behind eyes.Tubercles present behind eyes.In lateral view, angle formed by ventral margin of rostrum and head capsule c. 130°.Antennae.Scape reaching posterior margin of eyes when in repose, resting position running along ventral margin of rostrum.Funicle with 7 segments.Segments stout, clothed with dense thick setae; segments 1 and 2 lengths subequal, each about as long as 3 and 4 combined.Pronotum.Widest posteriorly, about 3/5 as wide as combined width of elytra; width at anterior margin much narrower than width at posterior margin.Lateral margins constricted in anterior 1/4 before abruptly widening, subparallel in posterior 2/3.Scales on disc larger than those on the elytra.Elytra.Stria 10 complete.Humeral callus developed.Disc without tubercles or spines.Wings.Reduced to fully developed.Costal margin straight, apex widely rounded.RA strongly sclerotised, and widest around middle.Radial cell (RC) completely sclerotised.RP 2 clearly evident.MP 1+2 wide and strongly sclerotised.CuA wide at base, divided in middle to form a long and narrow pseudocell.AP short, not reaching wing margin.Thoracic ventrites.Prosternum projecting ventrally, resulting in it having an anterior face.Mesoventral process swollen.Legs.All femora armed with a single ventral tooth; metafemoral tooth large, not excised at base of distal edge.All tibiae sinuous.Tarsal segment 1 stout, shorter than combined length of remaining segments; segment 5 about 1.5 times as long as segment 3. Male genitalia.Pedon tubular, relatively short, broad and high; membranous ventrally; base of pedon with a narrow, strongly sclerotised ventral brace.Temones approximately as long as pedon.Parameroid lobes elongate, fused along proximal 1/2.Manubrium stout, shorter than temones.Spiculum gastrale with furcal arms very broad, maximum width approximately 0.5 times length of apodeme.Female genitalia.Styli slender, inserted on ventral margin of gonocoxites.Gonocoxites short and broad.Bursa copulatrix long, apparently with two chambers.Sternite 8 entire, apex broadly rounded.Spermatheca C-shaped, slender.

Etymology
Named after Dr Guillermo Kuschel Gerdes (1918-2017) whose research into the weevils of the Southern Hemisphere gave substantial insight into weevil classification.The ending 'elysius' refers to Elysium, the 'Land of Joy' of Antiquity, which was located by Plato at the antipodes [32].

Biology
This species has frequently been collected on the flowerheads of Dracophyllum traversii Hook.f., and less commonly on Celmisia armstrongii Petrie.Further research is required to ascertain whether these records are an accurate indication of the larval host plant, or if the adults were only incidentally feeding on these plants.Adults have been collected at elevations between 1060 and 1520 m above sea level.Larvae are currently unknown.

Diagnosis
Evenly yellowish grey; small dark greyish brown maculae on elytra.Elytra long, 1.7 times longer than wide and 3.2 times longer than pronotum.Profemoral tooth small.

Biology
This species has been collected in leaf litter, in turf, on unidentified cushion plants and on Celmisia walkeri Kirk.A large series of specimens have been collected from Astelia nivicola Ckn.ex Cheesm.
The gut contents of the two dissected specimens collected from A. nivicola were filled almost exclusively with a single form of pollen.The specimen from Mt. Titiroa was collected in an unusual granite sand plain ecosystem [33,34].Adults have been collected at elevations between 900 and 1380 m above sea level.Larvae are currently unknown.

Etymology
Based on the Latin durus, 'strong, tough', in reference to the hardiness of this species which survives in the harsh environment of the Fiordland mountains.It is also a trait demonstrated by Willy, whose endurance while undertaking collecting expeditions in the Juan Fernandez Islands, New Zealand and elsewhere in the South Pacific is awe-inspiring.

Diagnosis
Mottled dark olive brown and greyish yellow.Mottling on elytra not forming any particular pattern, but forming paired dorsal vittae on the pronotum and broad bands on the distal 1/4 of the femora.Elytra relatively short, 1.5 times longer than wide and 3.2 times longer than pronotum.Profemoral tooth large (Figure 5g).

Etymology
Based on the Latin verbalis, 'of words,' an allusion to Willy's enjoyment of language.

Diagnosis
Uniformly medium grey; pronotum, elytra and legs with fine pale yellow-green setae that project over ground vestiture.Elytra long, 1.68 times longer than wide and 3.9 times longer than pronotum; declivity rounded in lateral view; elytral apices individually rounded.Profemoral tooth small.

Discussion
Although there have been several surveys of alpine insects in New Zealand [34,35], investigations into the evolutionary history of these taxa and the high level of endemism in these environments are still in their infancy [36].The alpine beetle fauna is quite rich, and almost every mountain range in the Southern Alps is home to endemic species [18,37].The New Zealand Eugnominae are common in alpine environments, with four other genera (Eugnomus Schönherr, 1847, Oreocalus May 1993, Pactolotypus Broun, 1909, Stephanorhynchus White, 1846) found in these areas [18].The description of this exclusively alpine weevil genus increases our understanding of the diversity of Coleoptera found in these habitats.
High-altitude eugnomine weevil richness and abundance may correlate with the high diversity of alpine plants.There are approximately 600 species of plants in the Southern Alps [38].The New Zealand alpine flora has an extraordinarily high proportion of white flowers, with the proportion of white flowered species in New Zealand being double that compared of other alpine regions in the world [39].It is thought that the pollinator fauna is rather depauperate and unspecialised [40], but the presence of eugnomine weevils and other beetles found exclusively on alpine flowers indicates that there is some level of host plant specialisation [41,42].Although beetles have been frequently acknowledged as being frequent flower visitors [39,40,43,44], they have not yet received specific attention to evaluate their role or effectiveness as pollinators in New Zealand ecosystems.Research in other countries has revealed a number of systems in which beetles, and weevils specifically, are primarily pollinators [45][46][47][48][49].It is possible that Kuschelysius and other alpine weevils may play an important role in pollination of alpine plants, including the pollination of high-altitude populations of the forest-inhabiting Dracophyllum traversii.
The difference in wing size between K. hollowayae and K. durus suggests that the importance of flight is very different for these two taxa.The shrubland and alpine forest habitat where K. hollowayae is found, is likely to require greater flight abilities than the tussock grassland and herbfields inhabited by K. durus, consistent with hypotheses that more homogenous environments promote flightlessness [50].Of these two, the larger species, K. hollowayae, was fully winged, the opposite of the general trend in New Zealand alpine stoneflies [51].Wing reduction or loss occurs as part of a syndrome of characters that repeatedly evolve in alpine insect taxa [52,53], like having a dark pigmented cuticle [37], or presence of quiescence instead of diapause [36].Flightlessness caused by wing reduction is likely to be one reason driving the species diversification in New Zealand alpine environments.In the Lucanidae, for example, wing reduction seems to have promoted speciation through the isolation of localised allopatric populations [54].
A full appreciation of the evolution of New Zealand's eugnomine weevil fauna, its relationships to host plants and the origin of the alpine fauna will require phylogenetic and faunistic studies coupled with more natural history observations.Furthermore, we expect to find additional species of Kuschelysius, especially in alpine areas with limited access.This paper is the first installment of what we hope will be a series of papers describing the New Zealand fauna and forming the basis for a full systematic treatment of the Eugnomini, which will underpin ecological and evolutionary studies.