Monograph |
Samuel D. J. Brown‡
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Corresponding author: Samuel D. J. Brown ( xsdjbx@gmail.com ) Academic editor: Miguel Alonso-Zarazaga
© 2017 Samuel D. J. Brown.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Brown SDJ (2017) Austromonticola, a new genus of broad-nosed weevil (Coleoptera, Curculionidae, Entiminae) from montane areas of New Zealand. ZooKeys 707: 73-130. https://doi.org/10.3897/zookeys.707.12649
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Abstract
Austromonticola gen. n. is proposed for a group of eight New Zealand alpine broad-nosed weevil species, all of which are here described: A. atriarius sp. n. (type locality: Umbrella Mountains, Central Otago), A. caelibatus sp. n. (type locality: Ohau Range, Mackenzie), A. furcatus sp. n. (type locality: Old Man Range, Central Otago), A. inflatus sp. n. (type locality: Hawkdun Range, Central Otago), A. planulatus sp. n. (type locality: St Marys Range, Central Otago), A. postinventus sp. n. (type locality: Kirkliston Range, South Canterbury), A. mataura sp. n. (type locality: Mt Dick, Otago Lakes) and A. rotundus sp. n. (type locality: Old Man Range, Central Otago). All species occur exclusively above 1000 m elevation in the mountains of Central Otago and South Canterbury in the South Island. A phylogeny of the genus, including six outgroups, was inferred from 33 morphological characters. It resolved the genus as monophyletic, and revealed two strongly supported clades within Austromonticola. DNA sequences of four gene regions were obtained from five species. Of these, the 3' end of COI proved to be the most suitable for the identification of specimens. Females of all species have diagnostic secondary sexual structures on the elytra and ventrites. These structures are hypothesised to have evolved to assist with oviposition in and beside cushion plants or by selection for structures to mitigate the costs to females of prolonged mating.
Keywords
Biodiversity, taxonomy, alpine, speciation, functional morphology
Introduction
The indigenous entimine weevil fauna of New Zealand currently consists of 28 described genera, containing 247 species. Taxonomic research on these weevils, especially at the genus level, has been dominated by the works of Francis Polkinghorne
The mountains of New Zealand are some of the most dramatic and recognisable landscapes of the country. Areas above 1000 m in elevation form a significant proportion of the available land area in the South Island. Geological evidence reveals that these landscapes have been formed relatively recently, with most ranges only appearing in the past five million years (
Materials and methods
Field collected specimens were killed in 100% ethanol or placed directly into a freezer at -20°C. Ethanol-preserved specimens were used preferentially for DNA extraction and sequencing.
Genitalia were examined by softening specimens for a short time in warm water, before removing the abdomen by inserting fine forceps between the metaventrite and ventrite 1. The abdomen was digested in porcine pancreatin enzyme solution for c. 36 h (
After clearing, the abdomen was flayed by cutting down the right side of the abdomen with spring scissors. Male genitalia were removed by severing the strong ligaments connecting sternite 8 to tergite 8, then cutting through the pretegminal membrane between the phallobase and the anus. Female genitalia were stained briefly by immersion in a 1g/l solution of Chlorazol Black in 70% ethanol, then removed by cutting through the membranes connecting tergites 7 and 8. Sternite 8 and tergite 8 were separated from the gonocoxites by cutting through their connecting membranes. Genitalia were photographed, then mounted on a card using dimethyl hydantoin formaldehyde (DMHF) (
Genitalia illustrations were prepared from photographs, using the program Inkscape (v. 0.91, Inkscape Team 2004-2017). Other line drawings were made with a Zeiss Stemi SV6 stereo microscope fitted with a camera lucida. These drawings were scanned and inked digitally in Inkscape. Habitus photographs were taken using a Nikon DS-Ri1 microscope fitted with a digital camera and a mechanical z-stepper. The program Nikon NIS Elements v. 4.10 was used to prepare the image stack and to produce the final montaged image.
Terminology follows
Descriptions of colour follow the terminology provided by the National Bureau of Standards (
Specimens were prepared for scanning electron microscopy (SEM) by separating the abdomen from the specimen, removing the tergites and genitalia and brushing down the sternites. Specimens were then air-dried before being mounted with double-sided carbon tape onto aluminium SEM stubs (11 mm high, 12 mm diameter). Specimens were coated with gold using a Emitech K975X sputter coater. Photographs were taken using a JEOL JSM-7000F field emission scanning electron microscope (JEOL, Tokyo, Japan), with an accelerating voltage of 3 kV.
Specimens were obtained and deposited in the following collections:
IACC Invermay Agricultural Centre Collection, Mosgiel, New Zealand
MONZ Te Papa Tongarewa, National Museum of New Zealand, Wellington, New Zealand
Label data of holotypes are transcribed using the following conventions. Data of individual labels are enclosed using quotes (‘…’), lines are indicated with a solidus (/) and metadata are given in square brackets ([…]).
Two-letter area codes follow the bioregionalisation system proposed by
A generative conception of species (
Data resources
Occurrence data from the specimens examined in this paper are deposited at GBIF, the Global Biodiversity Information Facility, http://ipt.pensoft.net/resource.do?r=austromonticoladistribution. DNA alignments and analysis scripts are available from FigShare, http://dx.doi.org/10.6084/m9.figshare.5367457
DNA sequencing and analysis
Only freshly collected specimens were used for sequencing. Genomic DNA was extracted from the pancreatin lysate (see above) using the Zymo Quick g-DNA Miniprep Kit (Zymo Research Corporation, Irvine, CA, U. S. A.), following the manufacturer’s instructions for a proteinase k extraction. Four gene regions were sequenced: the cytochrome c oxidase subunit I (COI) mitochondrial gene, the D2-D3 region of the 28S ribosomal RNA gene, the nuclear protein-coding gene arginine kinase (ArgK) and the nuclear protein-coding carbamoyl-phosphate synthetase 2-aspartate transcarbamylase-dihydroorotase (CAD) gene.
DNA was amplified using a 25 μl polymerase chain reaction (PCR) consisting of 1.25 U iStar Taq (iNtRON Biotechnology, Seongnam, South Korea), 0.4 mM dNTP, 1.5 mM MgCl2 and 0.2 μM of forward and reverse primers (Table
| Marker | Primer name | Direction | Primer sequence | Reference |
| COI | C1-J-2183 | Forward | 5'-CAA CAT TTA TTT TGA TTT TTT GG-3' |
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| LCO1490-JJ | Forward | 5'-CHA CWA AYC ATA AAG ATA TYG G-3' |
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| HCO2198-JJ | Reverse | 5'-AWA CTT CVG GRT GVC CAA ARA ATC A-3' |
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| TL2-N-3014 | Reverse | 5'-TCC AAT GCA CTA ATC TGC CAT ATT A-3' |
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| 28S | S3660 | Forward | 5'-GAG AGT TMA ASA GTA CGT GAA AC-3' |
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| 28S-Ff | Reverse | 5'-TTA CAC ACT CCT TAG CGG AT-3' |
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| ArgK | ArgKforB4 | Forward | 5'-GAY CCC ATC ATC GAR GAC TAC C-3' |
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| ArgKrevB1 | Reverse | 5'-TCN GTR AGR CCC ATW CGT CTC-3' |
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| CAD | CADfor4 | Forward | 5'-TGG AAR GAR GTB GAR TAC GAR GTG GTY CG-3' |
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| CADrev1mod | Reverse | 5'-GCC ATY RCY TCB CCY ACR CTY TTC AT-3' |
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Sequences were aligned by eye in Seaview (version 4.5.4) (
Morphological phylogenetic analysis
A total of 33 morphological characters were scored for 14 species (Table
The phylogenetic matrix was prepared using Mesquite (version 3.10) (
Taxonomic treatment
Austromonticola , gen. n.
Type species
Austromonticola mataura new species, here designated. Gender: masculine.
Diagnosis
Integument densely covered with small, grey appressed scales, elongate setiform scales ('setae') conspicuous along elytral interstriae. Rostrum stout, in dorsal view about 1.5 times longer than wide; subparallel proximally; scrobes lateral; ventral curvature with head capsule approximately 90°. Pronotum in dorsal view evenly convex. Elytra with small, shallow punctures, interstriae flat. Metanepisternal sutures complete. Metatibiae with apex simple. Penis tubular. Bursa copulatrix with a single sclerite.
Differential diagnosis
The combination of characters given above allows separation of Austromonticola from all other New Zealand weevils. The complete metanepisternal sutures distinguish them from Chalepistes Brown, 2017, in which the sutures are lacking. The abrupt 90° deflexion of the rostrum distinguish them from Catoptes Schönherr, 1842, which has a smoothly deflexed rostrum, angled about 120° with the ventral surface of the head capsule. The ridged appressed scales, conspicuous setae, evenly convex pronotum and small strial punctures separate Austromonticola from species of Inophloeus Pascoe, 1875 and Zenagraphus Broun, 1915, which have smooth appressed scales, inconspicuous setae, sculptured pronota and large, deep strial punctures. The subparallel rostrum and lateral scrobes, distinguish Austromonticola from Nicaeana Pascoe, 1877 and Haplolobus Broun, 1893, which have proximally widening rostra and dorsally situated scrobes.
Description
Body length ranging from 3.4 mm to 8.9 mm. Densely covered with appressed scales on all surfaces, interspersed with elongate setiform scales ('setae'); appressed scales on dorsum oval, 35–55 μm long, ridges visible at 30 × magnification, generally coloured bluish grey, brownish grey or blackish grey, easily abraded. Rostrum. Subparallel proximally in dorsal view, widened at antennal insertions. Epistome punctate, plurisetose, slightly raised above frons but separation indistinct. Epifrons with longitudinal median carina, lacking sulci; continuous with occiput, without distinct dorsal separation between head capsule and rostrum. Antennae. Sockets dorsolateral, situated in apical 1/3 of rostrum. Scapes clavate, reaching posterior margin of eye in repose. Funicular segments clavate, subspherical or oblately spheroid, moderately to loosely articulated, segments 7 almost as wide as club. Clubs two times longer than wide, tapering apicad. Head capsule. Interocular width in dorsal view greater than width of rostrum at base. Eyes large, lateral, flat, ovate to subcircular with long axis vertical, parallel with sagittal axis. Ventral curvature of head capsule and rostrum in lateral view angulate, approximately 90°. Pronotum. Disc in dorsal view smooth, evenly convex. Postocular lobes poorly to well developed; fringed with numerous short vibrissae attaining a maximum length of 1/3 times anterior-posterior length of eye. Elytra. In dorsal view approximately parallel-sided in anterior 2/3. Setae arising from interstriae. Elytral declivity in lateral view rounded in males, but sutural margin at top of declivity developed into tubercles in females of several species. Interstriae 3 above the declivity slightly swollen in both sexes of most species, interstriae 5 above the declivity rarely swollen. Ventral margin in lateral view sinuous, highest point near level of metacoxae. Thorax. Procoxae contiguous. Prosternum visible behind procoxae as a raised tubercle (“prosternellum”). Metaventrite with median suture visible only as a small, circular fovea posteriorly. Metanepisternal sutures complete. Abdomen. Ventrites 1 and 2 fused, subequal in length in middle; ventrites 3 and 4 subequal in length, approximately 0.5 times shorter than 1 or 2; ventrite 5 approximately equal in length to 1 or 2. Suture separating ventrites 1 and 2 curved anteriad in middle, other sutures straight. Wings. Absent. Legs. Uniformly and densely covered with appressed scales and setae, except for the posterior surface of the metafemora. Femora unarmed, maximum girth at about distal quarter. Pro- and mesotibiae with indistinct denticles along ventral margin and mucrones at apex; protibiae wider in distal 1/3 than proximal 1/3, incurved at apex. Metatibiae with dorsal and ventral margins subparallel; apical setal comb arcuate, pale; mucrones small, inconspicuous; without corbel. Tarsi with long, coarse setae on dorsal surface, without appressed scales; underside of segments 1 to 3 with short, dense setae forming pads medially divided by an inconspicuous glabrous line. Claws simple, separate, diverging. Male genitalia. Hemisternites 8 fully separate, with a forked membranous sclerite on the anterior margin of the membrane connecting them (‘spiculum relictum’,
Distribution
Restricted to alpine regions in Otago and South Canterbury, New Zealand.
Etymology
Derived from the Latin australis, meaning ‘southern’ and monticola, meaning ‘mountain dweller’, alluding to the habitat of the species of this genus, being confined to the mountains of the southern part of the South Island. Gender masculine.
Biology
Specimens of the genus have been collected in fellfield and cushionfield vegetation communities (Mark, 2012), commonly on top of, and close beside, cushion plants of the genera Phyllachne J. R. et G. Forst., 1776 (Stylidiaceae), Scleranthus L., 1753 (Caryophyllaceae), Veronica L., 1753 (Plantaginaceae), Hectorella Hook. f., 1864 (Montiaceae), Dracophyllum Labill., 1798 (Ericaceae) and Raoulia Hook. f., 1846 (Asteraceae), particularly when the plants have been in flower. Some species have also been found under specimens of Celmisia Cass., 1825 (Asteraceae) and Geum L., 1753 (Rosaceae). The larvae are as yet unknown.
Most specimens have been collected by hand collecting, though some have been captured in pitfall traps or by heat extraction from litter and turf samples.
Austromonticola atriarius , sp. n.
Diagnosis
Body size medium, 4 mm in length. Pronotum with median furrow. Elytral declivity in females with sutural tubercle; margin of ventrite 4 produced into a lamina, with bifurcate median process (Fig.
Description
Body length 3.60 mm to 4.55 mm (X‒ = 4.19 mm, s = 0.36, n = 8). Integument black. Dorsum densely covered with moderate yellowish brown to greyish brown appressed scales, pale “V” on elytral declivity often present; pronotum same colour as elytra, with pale posterolateral maculae lining up with pale maculae on humeral angles of elytra, especially prominent in females. Femora and tibiae with dense appressed scales concolorous with elytral scales, usually with pale band in distal 1/4 of femora. Tarsi with integument deep orange. Rostrum. Length 0.72 mm to 0.96 mm (X‒ = 0.85 mm, s = 0.09, n = 7), width 0.52 mm to 0.65 mm (X‒ = 0.59 mm, s = 0.05, n = 7), length/width ratio 1.33 to 1.55 (X‒ = 1.44, s = 0.07, n = 7). Epifrons with appressed scales imbricate; setae dolabriform, decumbent, dark; median and lateral carinae not evident. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with thick setae, without appressed scales. Antennae. Scapes in repose reaching hind margin of eye; covered with appressed scales and setae. Funicular segments moderately articulated; segments 1 clavate, subequal in length to 2; segments 2 clavate, about two times longer than 3; segments 3 to 4 clavate; segments 5 to 7 oblately spheroid. Pronotum. Length 1.07 mm to 1.29 mm (X‒ = 1.16 mm, s = 0.09, n = 7), width 1.58 mm to 2.08 mm (X‒ = 1.88 mm, s = 0.20, n = 7), length/width ratio 0.77 to 0.97 (X‒ = 0.87, s = 0.07, n = 7); in dorsal view widest in anterior 1/4, lateral margins evenly curved. Anterior margin slightly emarginate medially, posterior margin straight. Disc in dorsal view evenly convex, with median furrow; appressed scales imbricate; setae dolabriform to claviform, decumbent, dark to concolorous. Postocular lobes moderately developed. Elytra. Length 2.40 mm to 3.09 mm (X‒ = 2.78 mm, s = 0.26, n = 7), width 1.58 mm to 2.08 mm (X‒ = 1.88 mm, s = 0.20, n = 7), length/width ratio 1.39 to 1.61 (X‒ = 1.48, s = 0.08, n = 7). Anterior margin curved posteriad in middle, humeral angles rounded. Appressed scales imbricate. Setae claviform, decumbent, concolorous. Striae moderately impressed; interstriae slightly convex. Interstriae 1 at declivity flat in males, produced into a tubercle in females. Interstriae 3 and 5 at declivity swollen in both sexes. Apex in lateral view square in males; produced ventrad in females. Thoracic ventrites. Mesoventral process rounded. Mesanepisterna, mespimera, metanepisterna and metaventrite densely clothed with pappolepidia. Abdomen. Ventrites sparsely clothed, appressed scales most numerous medially, pappolepidia becoming dominant laterally. Apex rounded. Males with ventrite 1 strongly depressed medially; ventrite 5 flat. Females with ventrite 1 flat; ventrite 4 with posterior margin produced medially into a bifurcated lamina (Figs
DNA sequences. COI
KX191432. 28S. KX192009. ArgK. KX191719. CAD. KX191161.
Type material examined
Holotype. Female (
Paratypes. A total of 7 specimens (4 males, 3 females) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
CO. Gem Lake [45°34.236'S, 169°6.384'E, A], 14–15 Dec 1985, Barratt BIP, 1300 m (
Distribution
Fig.
Elevational range
Label data: 1300 m to 1430 m (X‒ = 1410 m, s = 46, n = 8). Georeferenced data: 1297 m to 1423 m (X‒ = 1313 m, s = 44, n = 8).
Etymology
From the Latin noun atriarius, ‘porter, doorkeeper’, in reference to the armature surrounding the female genital opening and alluding to a possible function of preventing unwanted mating attempts. The name is a noun in apposition.
Biology
Found in cushionfield, with a single specimen recorded in association with Phyllachne.
Austromonticola caelibatus , sp. n.
Diagnosis
Body size large, 8 mm in length. Epifrons flat, with semi-erect setae. Funicle segments 7 subconical. Pronotum evenly convex. Elytra with erect, piliform setae.
Description
Body length 7.83 mm to 8.91 mm (X‒ = 8.28 mm, s = 0.41, n = 5). Integument black. Dorsum densely covered with fine blackish blue appressed scales without metallic reflections. Femora and tibiae with appressed scales, unicolorous, concolorous with elytral scales. Tarsi with integument blackish red. Rostrum. Length 1.60 mm to 1.72 mm (X‒ = 1.65 mm, s = 0.05, n = 5), width 0.80 mm to 1.00 mm (X‒ = 0.94 mm, s = 0.08, n = 5), length/width ratio 1.68 to 2.04 (X‒ = 1.77, s = 0.15, n= 5). Epifrons with appressed scales tessellate; setae piliform, semi-erect, concolorous; median and lateral carinae evident. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae and appressed scales. Antennae. Scapes in repose reaching beyond hind margin of eyes; covered with appressed scales and setae. Funicular segments loosely articulated; segments 1 clavate, roughly as long as 2; segments 2 clavate, about two times longer than 3; segments 3 to 6 clavate, getting progressively shorter; segments 7 subconical. Pronotum. Length 1.90 mm to 2.24 mm (X‒ = 2.05 mm, s = 0.13, n = 5), width 3.08 mm to 3.25 mm (X‒ = 3.16 mm, s = 0.07, n = 5), length/width ratio 0.88 to 1.02 (X‒ = 0.94, s = 0.05, n = 5); in dorsal view widest in anterior 1/3, lateral margins evenly curved. Anterior margin entire, posterior margin straight. Disc in dorsal view evenly curved; appressed scales tessellate; setae piliform, semi-erect, concolorous. Postocular lobes moderately developed. Elytra. Length 5.15 mm to 5.61 mm (X‒ = 5.36 mm, s = 0.23, n = 5), width 3.08 mm to 3.25 mm (X‒ = 3.16 mm, s = 0.07, n = 5), length/width ratio 1.63 to 1.76 (X‒ = 1.70, s = 0.05, n = 5). Anterior margin slightly curved posteriad in middle, humeral angles rounded. Appressed scales tessellate. Setae piliform, semi-erect to erect, concolorous on disc, pale laterally and posteriorly. Striae moderately impressed; interstriae flat. Interstriae 1 at declivity flat in males; females unknown. Apex in lateral view square in males; females unknown. Thoracic ventrites. Mesoventral process rounded. Sutures between mesepimera and metanepisterna raised into a carina. Metaventrite densely covered with appressed scales. Abdomen. Ventrites sparsely clothed with appressed scales. Apex rounded. Males with ventrite 1 depressed medially, ventrite 5 flat. Females unknown. Male genitalia. Figs
DNA sequences
No DNA sequences obtained.
Type material examined
Holotype. Male (
Paratypes. A total of 4 specimens (4 males) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
MK: Lake Ohau Ski Field [44°13.44'S, 169°46.59'E, A], 15 Jan 2004, Johns PM, Nicholls D, 1600-1650 m (
Distribution
Fig.
Elevational range
Label data: 1625 m (n = 5). Georeferenced data: 1574 m (n = 5).
Etymology
From the Latin noun caelibatus, ‘celibacy’, an allusion to the fact that the species is thus far known only from the male sex; the species name is a noun.
Biology
No plant associations recorded.
Austromonticola furcatus , sp. n.
Diagnosis
Body size medium, 4 mm in length. Dense pappolepidia on venter. Elytral declivity in females with sutural tubercle; margin of ventrite 4 produced into a lamina, with deep median emargination (Fig.
Description
Body length 3.67 mm to 4.22 mm (X‒ = 3.98 mm, s = 0.19, n = 8). Integument black. Dorsum densely covered with brownish black to dark greyish yellowish brown appressed scales; pale “V” on elytral declivity largely confined to summits of protuberances; other pale variegations usually present on elytra, not forming patterns. Scutellum densely covered with pale scales. Pronotum same colour as elytra; with striking, pale, posterolateral maculae corresponding to pale maculae on humeral angles, especially in females. Femora and tibiae with appressed scales dense, unicolorous, concolorous with elytral scales. Tarsi integument dark reddish orange to black. Rostrum. Length 0.77 mm to 1.04 mm (X‒ = 0.90 mm, s = 0.09, n = 7), width 0.52 mm to 0.66 mm (X‒ = 0.57 mm, s = 0.05, n = 7), length/width ratio 1.35 to 1.68 (X‒ = 1.57, s = 0.11, n = 7). Epifrons with appressed scales imbricate; setae dolabriform, decumbent, concolorous; median and lateral carinae not evident. Dorsal carina arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae, without appressed scales. Antennae. Scapes in repose reaching hind margin of eyes; covered with appressed scales and setae. Funicular segments moderately articulated; segments 1 clavate, about 1.25 times longer than 2; segments 2 clavate, about 1.4 times longer than 3; segments 3 to 4 subspherical; segments 5 to 7 oblately spheroid, subequal in length. Pronotum. Length 1.06 mm to 1.24 mm (X‒ = 1.14 mm, s = 0.06, n = 7), width 1.61 mm to 2.16 mm (X‒ = 1.88 mm, s = 0.18, n = 7), length/width ratio 0.80 to 0.92 (X‒ = 0.86, s = 0.04, n = 7); in dorsal view widest in anterior 1/4, lateral margins strongly curved anteriorly, tapering posteriorly, more pronounced in females. Anterior margin slightly emarginate medially, posterior margin straight. Disc in dorsal view evenly curved; appressed scales imbricate; setae dolabriform, decumbent, dark to concolorous. Postocular lobes poorly developed. Elytra. Length 2.54 mm to 3.28 mm (X‒ = 2.78 mm, s = 0.25, n = 7), width 1.61 mm to 2.16 mm (X‒ = 1.88 mm, s = 0.18, n = 7), length/width ratio 1.39 to 1.58 (X‒ = 1.48, s = 0.07, n = 7). Anterior margin nearly straight, humeral angles rounded. Appressed scales tessellate to narrowly imbricate. Setae claviform, decumbent, pale to concolorous. Striae moderately impressed; interstriae flat on disc, convex on elytral declivity. Interstriae 1 at declivity flat in males, produced into a strong tubercle in females. Interstriae 3 and 5 at declivity swollen in both sexes. Apex in lateral view square in males; produced ventrad in females. Thoracic ventrites. Mesoventral process rounded. Metaventrite densely covered with pappolepidia. Abdomen. Ventrites clothed almost exclusively with pappolepidia, ventrites 1 and 2 moderately densely clothed, ventrites 3 to 5 increasingly sparse. Apex rounded. Males with ventrite 1 strongly depressed medially; ventrite 5 flat. Females with ventrite 1 flat; ventrite 4 with posterior margin produced into a subtriangular lamina with very deep apical emargination (Figs
DNA sequences. COI
KX191344. 28S. KX191914. ArgK. KX191626. CAD. KX191085.
Type material examined
Holotype. Female (
Paratypes. A total of 16 specimens (8 males, 8 females) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
CO: Hyde Rock [45°23.358'S, 169°11.844'E, A], 15 Mar 1975, Watt JC, 1524 m, Litter (
Distribution
Fig.
Elevational range
Label data: 1372 m to 1640 m (X‒ = 1509 m, s = 103, n = 16). Georeferenced data: 1583 m to 1665 m (X‒ = 1641 m, s = 19, n = 17).
Etymology
From the Latin adjective furcatus, ‘divided, forked’, in reference to the form of the ventral lamina of the female; the name is an adjective.
Biology
Specimens have been collected in association with Phyllachne cushions and Celmisia daisies. In particular, the largest series was associated with C. prorepens Petrie, 1887, but specimens have also been found with C. haastii Hook.f., 1864, and C. sessiliflora Hook.f., 1864.
Austromonticola inflatus , sp. n.
Diagnosis
Body size large, 8 mm in length. Rostrum with epifrons swollen (Fig.
Description
Body length 6.98 mm to 8.72 mm (X‒ = 7.94 mm, s = 0.64, n = 8). Integument black. Dorsum densely covered with fine appressed scales coloured greyish blue to dark greyish blue, often with brassy or purplish metallic reflections. Femora and tibiae with appressed scales unicolorous, concolorous with elytral scales. Tarsi with integument blackish red to dark red. Rostrum. Length 1.29 mm to 1.72 mm (X‒ = 1.60 mm, s = 0.15, n = 8), width 0.89 mm to 1.18 mm (X‒ = 1.05 mm, s = 0.09, n = 8), length/width ratio 1.37 to 1.64 (X‒ = 1.53, s = 0.09, n = 8). Epifrons swollen (Fig.
DNA sequences
COI. KX191461, KX191462. 28S. KX192043, KX192044, KX192045. ArgK. No sequences obtained. CAD. KX191187, KX191188.
Type material examined
Holotype. Female (
Paratypes. A total of 7 specimens (3 males, 4 females) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
CO: Hawkdun Range [44°47.256'S, 169°59.694'E, R], 11 Dec 2013, Brown SDJ, 1730 m, On Hectorella cushion (
Distribution
Fig.
Elevational range
Label data: 1680 m to 1730 m (X‒ = 1696 m, s = 23, n = 8). Georeferenced data: 1684 m to 1716 m (X‒ = 1699 m, s = 11, n = 8).
Etymology
From the Latin participle inflatus, ‘swollen, distended’, in reference to the convex epifrons of this species; the species name is a participle.
Biology
Specimens have been collected on Hectorella caespitosa Hook.f., 1864 and on and beside cushions of the snow hebe group (formerly placed in Chionohebe B.G.Briggs & Ehrend., 1976) of Veronica.
Austromonticola planulatus , sp. n.
Diagnosis
Body size large, 8 mm in length. Protibia with large denticles on ventral margin (Fig.
Description
Body length 7.59 mm to 8.25 mm (X‒ = 7.92 mm, s = 0.47, n = 2). Integument black. Dorsum covered with fine appressed scales, individual scales barely distinguishable, brownish black, with areas of brownish grey at sides of pronotum and base of rostrum. Femora and tibiae with appressed scales unicolorous, concolorous with elytral scales. Tarsi with integument black to strong red. Rostrum. Length 1.52 mm to 1.71 mm (X‒ = 1.62 mm, s = 0.13, n = 2), width 0.96 mm to 0.99 mm (X‒ = 0.98 mm, s = 0.02, n = 2), length/width ratio 1.58 to 1.73 (X‒ = 1.66, s = 0.10, n = 2). Epifrons with appressed scales imbricate; setae claviform, decumbent, concolorous; median and lateral carinae distinct, lateral carinae especially so. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae and appressed scales. Antennae. Scapes in repose reaching beyond hind margin of eyes; covered with appressed scales and setae. Funicular segments loosely articulated; segments 1 and 2 clavate, subequal, about 2 times longer than 3; segments 3 to 6 clavate, getting progressively shorter; segment 7 subconical. Pronotum. Length 1.92 mm to 2.28 mm (X‒ = 2.10 mm, s = 0.25, n = 2), width 3.22 mm to 3.47 mm (X‒ = 3.35 mm, s = 0.18, n = 2), length/width ratio 0.89 to 0.93 (X‒ = 0.91, s = 0.03, n = 2); in dorsal view widest in anterior 1/4, lateral margins evenly curved. Anterior margin sinuous, posterior margin straight. Disc in dorsal view evenly curved, except for median furrow extending from anterior 1/4 to posterior 1/8, deepest anteriorly; appressed scales imbricate; setae piliform to claviform, decumbent, dark. Postocular lobes strongly developed. Elytra. Length 5.21 mm to 5.39 mm (X‒ = 5.30 mm, s = 0.13, n = 2), width 3.22 mm to 3.47 mm (X‒ = 3.35 mm, s = 0.18, n = 2), length/width ratio 1.50 to 1.67 (X‒ = 1.59, s = 0.12, n = 2). Anterior margin almost straight, humeral angles rounded. Disc subdepressed. Appressed scales imbricate. Setae piliform to claviform, decumbent to semi-erect, concolorous on disc, pale laterally and posteriorly. Striae strongly impressed; interstriae convex; interstriae 1 at top of elytral declivity flat in males, swollen in females; interstriae 3 and 5 raised throughout length in both sexes. Apex in lateral view square in males; slightly produced ventrad and with small subapical tubercles in females. Thoracic ventrites. Mesoventral process narrowly rounded. Metaventrite densely covered with appressed scales. Abdomen. Ventrites densely covered with appressed scales. Males with ventrite 1 flat; ventrite 5 flat. Females with ventrite 1 flat; ventrite 4 with posterior margin produced laterally into small subtriangular laminae (Figs
DNA sequences
No DNA sequences obtained.
Type material examined
Holotype. Female (
Paratypes. A total of 1 specimen (1 male) designated as paratype, bearing blue paratype label. Paratype specimen deposited in
CO: Mt Bitterness [44°45.24'S, 170°18.198'E, A], 6–7 Feb 1978, Johns PM, Ingerfeld MH, 1830-1900 m, Stonefield with occasional mat plants (
Distribution
Fig.
Elevational range
Label data: 1865 m (n = 2).Georeferenced data: 1905 m (n = 2).
Etymology
From the Latin adjective planus, ‘flat, even’, combined with the the diminutive -ulus and the possessive -atus, referring to the almost level dorsum of this species, as compared with others in the genus; the species name is an adjective.
Biology
Collected in fellfield. No plant associations recorded.
Austromonticola postinventus , sp. n.
Diagnosis
Body size large, 8 mm in length. Epifrons flat, with decumbent setae. Females with ventrite 5 emarginate and with median swelling (Fig.
Description
Body length 7.26 mm to 8.42 mm (X‒ = 7.84 mm, s = 0.82, n = 2). Integument black. Dorsum densely covered with fine brownish black appressed scales with purple and gold metallic reflectance, reflectance particularly pronounced laterally and posteriorly. Femora and tibiae with appressed scales dense, unicolorous, concolorous with elytral scales. Tarsi with integument strong red. Rostrum. Length 1.45 mm to 1.67 mm (X‒ = 1.56 mm, s = 0.16, n = 2), width 0.90 mm to 1.02 mm (X‒= 0.96 mm, s = 0.08, n = 2), length/width ratio 1.61 to 1.64 (X‒ = 1.62, s = 0.02, n = 2). Epifrons with appressed scales tessellate; setae piliform, decumbent, pale; median and lateral carinae not evident. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae and appressed scales. Antennae. Fig.
DNA sequences
No DNA sequences obtained.
Type material examined
Holotype. Female (
Paratypes. A total of 1 specimen (1 male) designated as paratype, bearing blue paratype label. Paratype specimen deposited in
SC: Kirkliston Range [44°32.124'S, 170°30.954'E, A], 8–9 Feb 1978, Johns PM, Ingerfeld MH, 1740-1770 m, Stonefield with occasional mat plants (
Distribution
Fig.
Elevational range
Label data: 1755 m to 1829 m (n = 2). Georeferenced data: 1615 m to 1868 m (n = 2).
Etymology
From the Latin prefix post, ‘after’, and the participle inventus, ‘discovered’, referring to the recognition of this species after my PhD defence; the name is a participle.
Biology
Collected in fellfield. No plant associations recorded.
Austromonticola mataura , sp. n.
Diagnosis
Body size medium, 4 mm in length. Venter with glossy appressed scales, pappolepidia sparse. Elytral declivity of females with sutural tubercle; margin of ventrite 3 with paired median processes; margin of ventrite 4 produced into a lamina, with deep median emargination; margin of ventrite 5 with a broad horn on either side of the genital opening.
Description
Body length 3.42 mm to 4.11 mm (X‒ = 3.80 mm, s = 0.25, n = 13). Integument black. Dorsum densely covered with light bluish grey to dark greyish yellow appressed scales with metallic reflections; frequently with pale mottling sublaterally, and with scutellum, humeral area and hind pronotal angles bluish white; elytral declivity slightly paler than disc, especially in males. Pappolepidia of mesothoracic sternites light yellow. Femora appressed scales concolorous with elytral scales, often with obscure pale band in distal 1/4. Tarsi with integument deep orange to strong red. Rostrum. Length 0.72 mm to 0.88 mm (X‒ = 0.84 mm, s = 0.06, n = 8), width 0.52 mm to 0.66 mm (X‒ = 0.57 mm, s = 0.05, n = 7), length/width ratio 1.35 to 1.68 (X‒ = 1.57, s = 0.11, n = 7). Epifrons with appressed scales imbricate; setae claviform, decumbent, concolorous; median carina weak; lateral carinae evident. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae, without appressed scales. Antennae. Scapes in repose reaching middle of eyes. Funicular segments moderately articulated; segments 1 and 2 clavate, subequal, about 2 times longer than 3; segments 3 and 4 clavate, subequal; segments 5 to 7 subspherical, subequal. Pronotum. Length 1.06 mm to 1.18 mm (X‒ = 1.13 mm, s = 0.04, n = 8), width 1.61 mm to 2.16 mm (X‒ = 1.88 mm, s = 0.18, n = 7), length/width ratio 0.80 to 0.92 (X‒ = 0.86, s = 0.04, n = 7); in dorsal view widest in anterior 1/3, lateral margins strongly curved to widest point, gently curved behind. Anterior margin entire, posterior margin straight. Disc in dorsal view uneven, weak median furrow present, anterolateral depressions vague; appressed scales tessellate to imbricate; setae claviform, decumbent to semi-erect, concolorous. Postocular lobes poorly developed. Elytra. Length 2.30 mm to 3.04 mm (X‒ = 2.79 mm, s = 0.23, n = 8), width 1.61 mm to 2.16 mm (X‒ = 1.88 mm, s = 0.18, n = 7), length/width ratio 1.39 to 1.58 (X‒ = 1.48, s = 0.07, n = 7). Anterior margin curved posteriad in middle, humeral angles rounded. Appressed scales tessellate to imbricate. Setae claviform, decumbent to semi-erect, concolorous. Interstriae 1 at elytral declivity flat in males, produced into tubercles in females. Interstriae 3 raised at base; swollen at elytral declivity in both sexes, though more pronounced in females. Interstriae 5 at elytral declivity swollen in both sexes. Humeral region strongly pronounced by deeply impressed striae 9. Apex in lateral view square in males, produced ventrad in females. Thoracic ventrites. Mesoventral process narrowly rounded. Mesanepisterna, mesepimera and metanepisterna covered with small pappolepidia, contrasting with metaventrite densely covered with appressed scales. Abdomen. Ventrites clothed almost exclusively with appressed scales; ventrites 1 and 2 densely clothed in females, scales dense laterally and sparser medially in males; ventrites 3 to 5 increasingly sparse. Males with ventrite 1 depressed medially; ventrite 5 flat. Females with ventrite 1 flat; ventrite 4 posterior margin produced into a lamina, usually with a deep median emargination (Fig.
DNA sequences
COI. KX191347, KX191348, KX191349. 28S. KX191917, KX191918, KX191919. ArgK. KX191629, KX191630, KX191631. CAD. KX191088, KX191089, KX191090.
Type material examined
Holotype. Female (
Paratypes. A total of 21 specimens (9 males, 12 females) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
OL: Mt Dick [45°15.564'S, 168°40.926'E, R], 17 Jan 2014, Brown SDJ, 1600 m, On Dracophyllum muscoides cushion (
Distribution
Fig.
Elevational range
Label data: 1270 m to 1805 m (X‒ = 1580 m, s = 107, n = 23). Georeferenced data: 878 m to 1682 m (X‒ = 1479 m, s = 254, n = 23).
Etymology
This species is named after its distribution in the headwaters of the Mataura River; the name is a noun in apposition. The word mataura is Māori, of obscure meaning. Mataura was an ancestor of Ngatoro-i-rangi, the priest of the Arawa canoe. It may mean `glowing face', which is appropriate for its collection localities thus far have been on the eastern slopes of the Eyre Mountains.
Biology
Collected from Raoulia hectorii Hook.f., 1864, (recorded as R. haastii Hook.f., 1864, in error), moss, Dracophyllum muscoides Hook.f., 1864, and Phyllachne cushions. The majority of specimens were collected from Phyllachne.
Austromonticola rotundus , sp. n.
Diagnosis
Body size medium, 4.5 mm in length. Pronotum with subparallel lateral margins (Fig.
Description
Body length 4.20 mm to 4.80 mm (X‒ = 4.50 mm, s = 0.20, n = 11). Integument black. Dorsum densely covered with moderate olive to greyish brown appressed scales, some variegation usually present on elytra, but rarely forming distinct patterns; pronotum frequently with obscure lighter lines obliquely converging anteriorly. Femora and tibiae with dense appressed scales concolorous with elytral scales, usually with pale band in distal 1/4 of femur. Tarsi with integument deep orange. Rostrum. Length 0.89 mm to 0.99 mm (X‒ = 0.94 mm, s = 0.04, n = 6), width 0.58 mm to 0.66 mm (X‒ = 0.62 mm, s = 0.03, n = 6), length/width ratio 1.44 to 1.69 (X‒ = 1.52, s = 0.09, n = 6). Epifrons with appressed scales imbricate; setae claviform, decumbent, concolorous; median and lateral carinae not evident. Dorsal carinae arched over antennal insertions. Lateral area ventral of antennal insertions with fine setae and with appressed scales. Antennae. Scapes in repose reaching hind margin of eyes; covered with appressed scales and setae. Funicular segments moderately articulated; segments 1 clavate, about 1.5 times longer than 2; segments 2 clavate, about 1.2 times longer than 3; segments 3 and 4 clavate; segments 5 to 7 subspherical, subequal in length. Pronotum. Length 1.26 mm to 1.39 mm (X‒ = 1.32 mm, s = 0.05, n = 6), width 1.79 mm to 2.26 mm (X‒ = 1.98 mm, s = 0.19, n = 6), length/width ratio 0.83 to 0.91 (X‒ = 0.87, s = 0.03, n = 6); in dorsal view widest in anterior 1/3, lateral margins approximately subparallel (Fig.
DNA sequences
COI. KX191445. 28S. KX192022. ArgK. KX191729. CAD. KX191173.
Type material examined
Holotype. Female (
Paratypes. A total of 17 specimens (6 males, 11 females) designated as paratypes, bearing blue paratype label. Paratype specimens deposited in
CO: North Garvie Mountains, 9 Feb 1985, Barratt BIP, 1200 m, Ex Geum parviflorum (IACC: 1); Old Man Range [45°20.04'S, 169°12.534'E, A], 17 Jan 1965, Kuschel G, Townsend JI, 5000 feet (
Distribution
Fig.
Elevational range
Label data: 1200 m to 1590 m (X‒ = 1428 m, s = 76, n = 18). Georeferenced data: 1329 m to 1717 m (X‒ = 1477 m, s = 184, n = 17).
Etymology
From the Latin adjective rotundus, ‘round, spherical’ for the form of the female elytral declivity; the name is an adjective.
Biology
Specimens have been collected from Geum parviflorum Smith, 1805 and Dracophyllum muscoides. The majority of specimens, however, were collected from litter and turf (sward) samples.
Key to the species of Austromonticola
| 1 | Larger species, greater than 7 mm in length | 2 |
| – | Smaller species, less than 5 mm in length | 5 |
| 2(1) | Denticles on protibiae large, conspicuous (Fig. |
A. planulatus |
| – | Denticles on protibiae undeveloped (Fig. |
3 |
| 3(2) | Epifrons swollen, convex (Fig. |
A. inflatus |
| – | Epifrons flattened, level (Fig. |
4 |
| 4(3) | Epifrons with setae semi-erect. Setae along elytral interstriae 7 erect | A. caelibatus |
| – | Epifrons with setae decumbent. Setae along elytral interstriae 7 decumbent | A. postinventus |
| 5(1) | Pronotum hexagonal in outline, widest anteriorly, sides evenly converging toward base (Fig. |
6 |
| – | Pronotum round in outline (Fig. |
A. rotundus |
| 6(5) | Venter with dense pappolepidia, round appressed scales sparsely distributed | 7 |
| – | Venter with dense appressed scales, pappolepidia sparsely distributed | A. mataura |
| 7(6) | Pronotum with median furrow. Elongate elytral scales decumbent. Antennal funicle segments 3 longer than 4 | A. atriarius |
| – | Pronotum evenly curved. Elongate elytral scales semi-erect. Antennal funicle segments 3 of similar length as 4 | A. furcatus |
Molecular diagnostics
Specimens of five species of Austromonticola were available for DNA sequencing. No fresh specimens of A. planulatus, A. caelibatus and A. postinventus were collected. Multiple specimens were available only of A. inflatus and A. mataura, and only the latter yielded multiple sequences for all gene regions. Due to these low sample numbers, conclusions regarding intra-specific variability are necessarily limited.
The three protein-coding genes could all be unambiguously aligned, 28S being the only locus that required alignment gaps. The COI alignment was divided into two regions. The first represented the 5' region, corresponding to the region favoured for DNA barcoding (
Genetic variation existed in all gene regions, COI showing the greatest amount of variation, followed by CAD and ArgK, and 28S displaying the least (Figs
COI proved to be the most suitable gene for identifying specimens of Austromonticola. The 3' end of COI allowed unambiguous differentiation of all species with available data. This region has the greatest taxon coverage, though indications are that the 5' ‘barcoding’ end of the gene has higher levels of variation if amplification is successful (Fig.
The same pattern of variation in each gene region was observed when the number of diagnostic nucleotides was calculated (Figs
Across all three protein-coding genes, A. atriarius, A. furcatus and A. mataura showed the smallest interspecific distances. In COI, A. rotundus was nearest to A. inflatus (Fig.
Phylogenetic analysis
Analysis of the character matrix (Table
Character matrix for cladistic analysis of relationships within Austromonticola.
| Taxon | 1 | 6 | 11 | 16 | 21 | 26 | 31 |
|---|---|---|---|---|---|---|---|
| Austromonticola atriarius | 02100 | 00100 | 11000 | 11011 | 10010 | 01101 | 11? |
| Austromonticola caelibatus | 12101 | 00010 | 00001 | 1?21? | 0???? | ???01 | 021 |
| Austromonticola furcatus | 02101 | 00100 | 11000 | 01011 | 10010 | 11101 | 111 |
| Austromonticola inflatus | 12101 | 00010 | 00001 | 01210 | 00110 | 11101 | 021 |
| Austromonticola planulatus | 12100 | 00000 | 00000 | 10211 | 01010 | 01111 | 011 |
| Austromonticola postinventus | 12101 | 00010 | 00001 | 01210 | 00110 | 01101 | 021 |
| Austromonticola mataura | 02101 | 00100 | 11000 | 11011 | 10010 | 11101 | 111 |
| Austromonticola rotundus | 02101 | 00100 | 00000 | 10011 | 00010 | 01111 | 01? |
| Irenimus parilis | 00110 | 00001 | 00000 | 10000 | 01021 | 00020 | 000 |
| Brachyolus punctatus | 00010 | 01000 | 00000 | 00100 | 01000 | 00000 | 010 |
| Inophloeus sulcifer | 11001 | 01000 | 01110 | 00100 | 01011 | 00001 | 011 |
| Zenagraphus metallescens | 11000 | 11000 | 00110 | 00000 | 01011 | 02100 | 010 |
| Inophloeus sternalis | 10000 | 01001 | 00000 | 10100 | 00021 | 00001 | 021 |
| Undescribed genus and species | 11000 | 10000 | 1001? | 00100 | 00010 | 00120 | 021 |
In the discussion of characters below, the significance of synapomorphies is only discussed in relation to Austromonticola, due to limited representation of outgroup taxa.
1. Body length, as measured from the anterior margin of the eyes to the elytral declivity in lateral view: (0) less than 6 mm; (1) greater than 6 mm. As estimated by this phylogeny, state 1 is the ancestral body size, while state 0 is homoplasious for the A. rotundus–A. mataura clade and Irenimus parilis + Brachyolus punctatus (ci = 0.5, ri = 0.8).
2. Body vestiture, form of appressed scales covering dorsum: (0) scales large, conspicuous, imbricate, coloured brown or bronze or pale yellow; (1) scales small, inconspicuous, tessellate, dome-shaped, black, ridges not visible at 50 × magnification; (2) scales small, tessellate, flat, coloured greys or browns, ridges visible at 30 × magnification. State 2 is a synapomorphy for Austromonticola (ci = 1, ri = 1).
3. Labium, form of base: (0) flat; (1) concave, with lateral areas raised relative to disc. State 1 is a synapomorphy for Austromonticola but convergently present in Irenimus parilis (ci = 0.5, ri = 0.75).
4. Labium, setation of disc: (0) bare; (1) with setae distally and laterally. The plesiomorphic state 0 is present in all species of Austromonticola, with state 1 being a synapomorphy for Irenimus parilis + Brachyolus punctatus (ci = 1, ri = 1).
5. Rostrum, ventral side, hypostomal-labial sutures: (0) strongly convergent to point distal of head capsule deflexion; (1) roughly parallel, converging to point proximal of head deflexion. State 1 is convergently present in Inophloeus sulcifer and Austromonticola excluding A. planulatus, though with a reversal in A. atriarius (ci = 0.33, ri = 0.67).
6. Frons, vestiture: (0) clothed with pale, thick, conspicuous setae; (1) clothed with inconspicuous setae. The plesiomorphic state 0 is present in all species of Austromonticola, with state 1 being a synapomorphy for Zenagraphus metallescens + the undescribed genus (ci = 1, ri = 1).
7. Pronotum, sculpture of disc: (0) evenly convex, without obvious sculpture; (1) with depressions or wrinkles laterally. State 0 is shared with the undescribed genus and I. parilis by all species of Austromonticola (ci = 0.33, ri = 0.33).
8. Pronotum, scale pattern: (0) without distinctive pattern; (1) with lateral vittae formed by pale scales, vittae extending onto humeral area. State 1 is a synapomorphy for the A. rotundus–A. mataura clade (ci = 1, ri = 1).
9. Protarsus, second segment: (0) transverse, or width subequal to length; (1) distinctly elongate, length greater than width. State 1 is a synapomorphy for the A. caelibatus–A. inflatus clade (ci = 1, ri = 1).
10. Metatibial apex: (0) simple, without corbel; (1) with bare corbel. The plesiomorphic state 0 is present in all species of Austromonticola. This phylogeny estimates that state 1 is convergently present in Inophloeus sternalis and Irenimus parilis (ci = 0.5, ri = 0).
11. Metanepisterna, vestiture: (0) consisting of round appressed scales; (1) consisting of pappolepidia. State 1 is a synapomorphy for the A. atriarius–A. mataura clade, but convergently present in the undescribed genus (ci = 0.5, ri = 0.67).
12. Metanepisterna, vestiture: (0) comprising two rows of scales; (1) comprising three or more rows of scales. State 1 is a synapomorphy for the A. atriarius–A. mataura clade but convergently present in I. sulcifer (ci = 0.5, ri = 0.67) (ci = 0.5, ri = 0.67).
13. Elytra, form of strial punctures: (0) shallow, circular, interstriae much wider than punctures; (1) deep, subquadrate, interstriae approximately equal in width as punctures. The plesiomorphic state 0 is present in all species of Austromonticola. State 1 is convergently present in Inophloeus sulcifer and Zenagraphus metallescens (ci = 0.5, ri = 0).
14. Elytra, length and density of setae on disc: (0) long, conspicuous and evenly distributed; (1) short, sparse. The plesiomorphic state 0 is present in all species of Austromonticola. State 1 is a synapomorphy for the Inophloeus sulcifer–Zenagraphus metallescens clade (ci = 1, ri = 1).
15. Elytra, form of setae: (0) claviform; (1) piliform. State 1 is a synapomorphy for the A. caelibatus–A. inflatus clade (ci = 1, ri = 1).
16. Elytra, humeral region anteriorly of conjunction of striae 7 & 8: (0) evenly convex, not prominent in comparison with surroundings, stria 9 not deeply impressed at base; (1) strongly raised in comparison with surroundings, stria 9 deeply impressed at base. A complex character that does not show any clear relationships (ci = 0.25, ri = 0.5).
17. Elytra, form of sutural interval above elytral declivity in females: (0) flat; (1) developed into a prominent tubercle. State 1 is convergently present in the A. caelibatus–A. inflatus clade and the A. atriarius–A. mataura clades (ci = 0.5, ri = 0.75). The unknown female of A. caelibatus is predicted by this estimation to possess state 1.
18. Elytra, form of interstriae 3 above elytral declivity: (0) swollen, more so than on disc; (1) developed into prominent tubercles; (2) flat, no greater than on disc. State 2 is shared between A. planulatus and the A. caelibatus–A. inflatus clade. State 0 is a shared by all members of the A. rotundus–A. mataura clade but has also evolved elsewhere in the tree (ci = 0.5, ri = 0.67).
19. Elytra, form of interstriae near apex: (0) clearly impressed, confluence of interstriae 3 and 9 clearly raised above confluence of 2 and 10; (1) striae obsolete, confluence of interstriae 3 and 9 on same level as confluence of 2 and 10. State 1 is a synapomorphy for Austromonticola (ci = 1, ri = 1).
20. Ventrite 4 of females, form of posterior margin: (0) entire; (1) produced into a lamina. State 1 is a synapomorphy for Austromonticola, but a reversal to state 0 has occurred in A. postinventus and A. inflatus (ci = 0.5, ri = 0.75).
21. Ventrite 5 of females, form of apex: (0) entire or only slightly emarginate; (1) strongly emarginate, flanked by horns. State 1 is a synapomorphy for the A. atriarius–A. mataura clade (ci = 1, ri = 1).
22. Female genitalia, presence of rectal valve (
23. Female genitalia, length/height ratio of distal gonocoxite: (0) long and slender, ratio greater than 1.8; (1) stout, ratio less than 1.5. State 1 is a synapomorphy for the A. caelibatus–A. inflatus clade (ci = 1, ri = 1). The unknown female of A. caelibatus is predicted by this phylogeny to possess state 1.
24. Female genitalia, number of sclerites in the bursa copulatrix: (0) absent; (1) one present; (2) two present. This cladogram infers that a single bursal sclerite is the plesiomorphic state, which is shared by all species of Austromonticola. State 0 is an apomorphy for Brachyolus punctatus, while state 2 is convergently present in Inophloeus sternalis and Irenimus parilis (ci = 1, ri = 1).
25. Female genitalia, form of vagina: (0) unsclerotised; (1) sclerotised. State 1 is a character uniting all species of Austromonticola but shared with Brachyolus punctatus and the undescribed genus (ci = 0.33, ri = 0.33).
26. Female genitalia, caudal shape of sclerotised rods on proximal gonocoxite: (0) straight in ventral view; (1) curved inwardly in ventral view. State 1 has arisen twice, in A. furcatus–A. mataura and apparently independently in A. inflatus (ci = 0.5, ri = 0.5).
27. Female genitalia, shape of sclerotised rods on proximal gonocoxite: (0) tapering proximally; (1) broadening proximally; (2) strongly broadening proximally, multiply divided. State 1 is a synapomorphy for Austromonticola (ci = 1, ri = 1).
28. Female genitalia, position of sclerotised rods on proximal gonocoxite in lateral view: (0) median; (1) ventral. State 1 unites all species of Austromonticola, but is shared with Zenagraphus metallescens and the undescribed genus (ci = 0.5, ri = 0.67).
29. Penis, apex in dorsal view: (0) acute; (1) sagittate; (2) truncate. State 0 is the usual form in Austromonticola, but state 1 has arisen twice, in A. planulatus and A. rotundus (ci = 0.5, ri = 0).
30. Penis, curvature in lateral view: (0) even from base to apex, maximum height near middle; (1) largely confined to base, maximum height near basal 1/3. The plesiomorphic state 1 is present in all species of Austromonticola. State 0 is a homoplasious synapomorphy for Brachyolus punctatus + Irenimus parilis and Zenagraphus metallescens + the undescribed genus (ci = 0.5, ri = 0.67).
31. Penis, form of ostial region: (0) tubular, unmodified; (1) thickened to form sclerotised crest (Fig.
32. Penis, length of temone in relation to length of pedon in lateral view: (0) longer than pedon; (1) shorter than pedon, but longer than 0.7 times length of pedon; (2) shorter than 0.7 times length of pedon. State 2 is a homoplasious character that unites the members of the A. caelibatus–A. inflatus clade but shared with the undescribed genus (ci = 0.5, ri = 0.5).
33. Male genitalia, shape of hemisternites 8: (0) roughly quadrate; (1) roughly triangular. The plesiomorphic state 1 is present in all species of Austromonticola. State 0 unites Brachyolus punctatus + Irenimus parilis, but is convergently present in Zenagraphus metallescens (ci = 0.5, ri = 0.5).
The species of Austromonticola are united by three unambiguous synapomorphies, scale structure (character 2), the obsolete striae at the elytral apex (character 19), and the proximally widening gonocoxal rods (character 27).
In Austromonticola there are two strongly supported clades, one consisting of the larger species A. caelibatus, A. postinventus and A. inflatus (the A. inflatus clade) and the other consisting of the smaller species possessing metanepisterna with three rows of pappolepidia, a penis with an ostial crest and ventrite 5 with a strongly emarginate apex in the females, A. atriarius, A. furcatus and A. mataura (the A. mataura clade). Grouped with the latter clade is A. rotundus; however, the support for this relationship is weak, and the sagittate apex of the penis shared with A. planulatus hints at a possible relationship. Together, A. planulatus and A. rotundus provide a transitional series between the distinctly different A. inflatus clade and the A. mataura clade.
Discussion
Development of ventrites in females
The highly modified ventrites in many species of Austromonticola are a particularly fascinating feature of the genus. There is a range of developments of the posterior margin of ventrite 4 of the females. No laminae are found in A. inflatus and A. postinventus, rather the posterior margin of ventrite 4 is recurved anteriad. A short lamina with a wide emargination is present in A. planulatus. Long bifurcate laminae are found in A. mataura and A. atriarius, while A. furcatus has a broader lamina with a deep emargination. Finally, A. rotundus has a long, broad lamina with a shallow emargination. The unknown female of A. caelibatus is predicted in the phylogenetic tree inferred above to be lacking a lamina, but it is equally parsimonious to infer a lamina being present in A. caelibatus, given the basal position of the species in the clade. The form of the lamina in this species would be of interest. This range of development make Austromonticola a suitable system for investigating the function of these laminae. Two hypotheses are presented in detail here.
Preparation of oviposition sites
The first hypothesis is that these ventral structures assist in the preparation of oviposition sites in cushion plants. The cushion vegetational form is a distinctive feature of New Zealand alpine plants, such as Raoulia and Phyllachne. This growth habit presents densely packed foliage underlain by a peaty layer formed by decaying leaves still attached to the plant (
This hypothesis also provides an explanation for the recurved margin of ventrite 4 of A. inflatus and A. postinventus. In these species, it is hypothesised that the form of ventrite 4 allows maximum flexion of ventrite 5, which assists in ovipositing under the side of the cushion plants where the plant meets the surrounding substrate (Fig.
The rather different laminae of A. planulatus and A. rotundus suggest different oviposition behaviours or host plants from those of the two scenarios postulated above.
Mate hindrance
The second hypothesis is that these structures are mate hindrance devices. Mating pairs of entimine weevils are frequently encountered in copula in the field, and studies of their mating behaviour in captivity show that males will remain mounted on females for extended periods of time (D Watkin and SDJ Brown, unpub. data). The costs imposed by extended mounting include the energy expended in carrying males (
The two hypotheses presented above are not necessarily mutually exclusive. These two selection pressures may act synergistically, which may explain the rapid evolution of these structures. Further observations of oviposition and mating behaviour of Austromonticola, combined with experiments manipulating the form of the laminae, will be required to evaluate these hypotheses.
Other, alternate hypotheses for these ventral structures could include male stimulation during mating, pre-copulatory species recognition signals to prevent hybridisation, assisting the retraction of the ovipositor after oviposition has been completed and providing an area for sensory organs to determine optimum oviposition sites.
Modified ventrites in other weevils
Although unusual, the highly modified ventrites of Austromonticola females are not unique. In the New Zealand context, modified ventrites are also known in species of Chalepistes (e.g. C. dugdalei (Barratt & Kuschel, 1996), C. curvus (Barratt & Kuschel, 1996) and C. patricki (Barratt & Kuschel, 1996)) and Nicaeana, which have medial laminae on ventrite 4 or various swellings on ventrite 5 (
The cushion growth form is a feature of alpine vegetation worldwide, and is prevalent in the Himalayas (
Relationships
The morphological phylogeny is largely consistent with the molecular data, in that both indicate a close relationship between A. furcatus, A. atriarius and A. mataura. However, the position of A. rotundus in the morphology-based tree, placed as the sister taxon of the A. mataura clade, is not supported by the molecular data. The overall signal from the molecular data is that A. rotundus is the most distant of all the species for which DNA sequences were obtained, however no consensus was gained regarding its nearest relative. Results from COI and 28S are surprising. In the analysis of COI, A. inflatus was nearest to A. rotundus, whereas A. inflatus has the same 28S sequence as A. atriarius and A. furcatus. These results serve to bolster confidence that A. inflatus, A. postinventus and A. caelibatus are congeneric with the other members of the genus. Obtaining DNA sequences from the other species in the genus, especially the morphologically distinct A. planulatus, will be important for further insights into the relationships of species in the genus.
New Zealand alpine weevil fauna
Austromonticola is one of a number of weevils that inhabit the montane regions of New Zealand. Other weevil genera with representatives found above the treeline include Baeosomus Broun, 1904 (Brachycerinae), Anagotus Sharp, 1882, Gromilus Blanchard, 1853, Liparogetus Broun, 1915, Lyperopais Broun, 1893 (Cyclominae), Lyperobius Pascoe, 1876, “Crisius” Pascoe, 1876 (Molytinae), Eugnomus Schönherr, 1847, Oreocalus May 1993, Pactolotypus Broun, 1909, Stephanorhynchus White, 1846 (Curculioninae: Eugnomini), Peristoreus and Simachus (Curculioninae: Storeini). However, the Entiminae are best represented, with 13 genera (Austromonticola, Sargon Broun, 1903, Inophloeus, Chalepistes, Catoptes, Nicaeana, Haplolobus, Zenagraphus, Neoevas Broun, 1921, and four undescribed genera) having species found primarily or solely in montane environments over 1000 m in elevation.
This diverse community is at apparent odds with the young geological age of the environment. The Southern Alps began rising around 5 million years ago (
A geobiological model of the origin of the New Zealand alpine flora posited by
An alternative possibility for the origin of the New Zealand alpine biota is dispersal from alpine regions in Australia, South America or the Northern Hemisphere. These areas have been the main sources for the majority of New Zealand alpine plant radiations (
Research into the mechanisms by which these weevil lineages have adapted to alpine environments, as has been investigated in other New Zealand alpine insects (
Conservation significance
The localised distribution of most of the species of Austromonticola place them within the Naturally Uncommon (Range Restricted) threat classification category (
Conclusion
Additional research into the biology, behaviour and physiology of the species of Austromonticola described here will offer insight into the function of the exaggerated abdominal structures of the females, and into processes by which sexual selection accelerates speciation. Further exploration and collecting, especially in areas such as the Mt Teviot/Manorburn region in Central Otago, the Pisa Range, Dunstan Mountains and Mt Aspiring National Park, will be vital for discovering additional species in the genus, which will provide further data for evaluating hypotheses of the role of historical contingency and environmental pressures on the evolution of alpine insects.
Plates
Genitalia of Austromonticola atriarius. 39 penis, dorsal view 40 aedeagus, lateral view 41 female tergite 8, dorsal view 42 ovipositor, dorsal view 43 ovipositor and spermatheca, lateral view 44 bursal sclerite, ventral view 45 female sternite 8, ventral view. Scale bars = 0.5 mm; 39–40 at same scale; 41–45 at same scale.
Genitalia of Austromonticola caelibatus. 46 penis, dorsal view 47 aedeagus, lateral view 48 male hemisternites 8 and spiculum gastrale, lateral view (membrane between hemisternites 8 and basal plate indicated) 49 male hemisternites 8 and spiculum gastrale with basal plate, ventral view. Scale bar = 0.5 mm.
Genitalia of Austromonticola furcatus. 50 penis, dorsal view 51 aedeagus, lateral view 52 male hemisternites 8 and spiculum gastrale, lateral view (membrane between hemisternites 8 and basal plate indicated) 53 male hemisternites 8 and spiculum gastrale with basal plate, ventral view 54 female tergite 8, dorsal view 55 ovipositor, dorsal view 56 ovipositor and spermatheca, lateral view 57 female sternite 8, ventral view. Scale bars = 0.5 mm; 50–53 at same scale; 54–57 at same scale.
Genitalia of Austromonticola inflatus. 58 penis, dorsal view 59 aedeagus, lateral view 60 male hemisternites 8 and spiculum gastrale, lateral view (muscles between hemisternites 8 and basal plate indicated) 61 male hemisternites 8 and spiculum gastrale with basal plate, ventral view 62 female tergite 8, dorsal view 63 ovipositor, dorsal view 64 bursal sclerite, anterior view 65 ovipositor and spermatheca, lateral view 66 sternite 8, ventral view. Scale bars = 0.5 mm; 58–61 at same scale; 62–66 at same scale.
Genitalia of Austromonticola planulatus. 67 penis, dorsal view 68 aedeagus, lateral view 69 male hemisternites 8 and spiculum gastrale, lateral view (muscles between hemisternites 8 and basal plate indicated) 70 male hemisternites 8 and spiculum gastrale with basal plate, ventral view 71 tergite 8, dorsal view 72 ovipositor, dorsal view 73 bursal sclerite, anterior view 74 ovipositor and spermatheca, lateral view 75 female sternite 8, ventral view. Scale bars = 0.5 mm; 67–70 at same scale; 71–75 at same scale.
Genitalia of Austromonticola postinventus. 76 penis, dorsal view 77 aedeagus, lateral view 78 male hemisternites 8 and spiculum gastrale, lateral view (muscles between hemisternites 8 and basal plate indicated) 79 male hemisternites 8 and spiculum gastrale with basal plate, ventral view 80 female tergite 8, dorsal view 81 ovipositor, dorsal view 82 bursal sclerite, anterior view 83 ovipositor and spermatheca, lateral view 84 female sternite 8, ventral view. Scale bars = 0.5 mm; 75–79 at same scale; 80–84 at same scale.
Genitalia of Austromonticola mataura. 85 aedeagus, dorsal view 86 aedeagus, lateral view 87 male hemisternites 8 and spiculum gastrale, lateral view (muscles between hemisternites 8 and basal plate indicated) 88 male hemisternites 8 and spiculum gastrale with basal plate, ventral view 89 female tergite 8, dorsal view 90 ovipositor, dorsal view 91 ovipositor and spermatheca, lateral view 92 female sternite 8, ventral view. Scale bars = 0.5 mm; 85–88 at same scale; 89–92 at same scale.
Genitalia of Austromonticola rotundus. 93 penis, dorsal view 94 aedeagus, lateral view 95 female tergite 8, dorsal view 96 ovipositor, dorsal view 97 ovipositor and spermatheca, lateral view 98 bursal sclerite, ventral view 99 female sternite 8, ventral view. Scale bars = 0.5 mm; 93–94 at same scale; 95–99 at same scale.
Cladograms showing phylogenetic relationships between Austromonticola species as inferred from morphological data. 118 single most parsimonious tree inferred from 33 characters scored for 14 species 119 phylogenetic tree of Austromonticola with bootstrap values above nodes and jackknife values below. Nodes with lower than 50% support were collapsed.
Heatmaps of the uncorrected pairwise genetic distances between Austromonticola specimens sampled. Lighter colours indicate greater distances. 120 the 3' region of the COI mitochondrial protein-coding gene 121CAD nuclear protein-coding gene 122 the 5' (“barcoding”) region of COI123ArgK nuclear protein-coding gene 124 28S nuclear ribosomal RNA gene.
Diagnostic nucleotides within the Austromonticola specimens sampled. Numbers above the bars indicate the position of the diagnostic base within the alignment. Numbers in parentheses beside species names indicate the numbers of specimens included in the alignment. Letters below the bars and the colour of the vertical bar indicate the value of the diagnostic nucleotide. 125 the 5' (“barcoding”) region of the COI mitochondrial protein-coding gene 126 the 3' region of COI127CAD nuclear protein-coding gene 128ArgK nuclear protein-coding gene 129 28S nuclear ribosomal RNA gene.
Schematic diagrams of hypothesised oviposition posture. 130 hypothesised function of lamina on ventrite 4 and horns surrounding genital orifice on ventrite 5, which force apart dense foliage of cushion plants to allow oviposition in peaty layer underneath 131 hypothesised function of recurved margin of ventrite 4 which allows maximum flexion of ventrite 5, enabling oviposition under the side of cushions between the plant and the surrounding substrate. Abbreviations: V4–ventrite 4; V5–ventrite 5. Figures not drawn to scale.
Acknowledgements
This research was funded by Lincoln University, the Miss E. L. Hellaby Indigenous Grasslands Research Trust and AgResearch (MSI contract LINX0304, Ecosystems Bioprotection), as part of the author’s doctoral programme. The basemap of distribution maps was provided courtesy of Geographx. Miguel Alonso-Zarazaga (Museo Nacional de Ciencias Naturales), Robert Anderson (Canadian Museum of Nature), Karen Armstrong (Bio-Protection Research Centre), John Marris (Bio-Protection Research Centre) and Rolf Oberprieler (CSIRO Ecosystem Services) critically commented on the manuscript. Robert Hoare (Landcare Research) provided guidance on name formation. Mike Flaws (University of Canterbury) assisted with scanning electron microscopy, while Cor Vink (Canterbury Museum) provided access to photomicroscopy facilities. Gregory Holwell (University of Auckland), Ren Li (Institute of Zoology, Chinese Academy of Sciences) and Greg Setliff (Kutztown University) provided useful comments and discussion about morphological features.
The following curators and institutions that provided specimens are most gratefully acknowledged: Barbara Barratt (Invermay Agricultural Centre Collection, Mosgiel), Rich Leschen (New Zealand Arthropod Collection, Auckland), John Marris (Lincoln University Entomology Research Museum, Christchurch), and Cor Vink (Canterbury Museum, Christchurch).
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