Zootaxa 4263 (1): 001–042 ISSN 1175-5326 (print edition) Article ZOOTAXA http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4263.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:78E0852E-F66A-44E4-8798-E5187185C950 A revision of the New Zealand weevil genus Irenimus Pascoe, 1876 (Coleoptera: Curculionidae: Entiminae) SAMUEL D. J. BROWN1,2,3 1Bio-Protection Research Centre, PO Box 85084, Lincoln University 7647, Canterbury, New Zealand 2AgResearch, Gerald St, Lincoln, Canterbury, New Zealand 3Correspondence. E-mail: [email protected] Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Irenimus Pascoe, 1876. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Key to species of Irenimus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 1. Irenimus aniptus Brown, new species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 2. Irenimus crinitus Brown, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 3. Irenimus duplex (Broun, 1904) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 4. Irenimus minimus Brown, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 5. Irenimus parilis Pascoe, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 6. Irenimus stichus Brown, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 7. Irenimus thoracicus Brown, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24 Molecular diagnostics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27 Changes in the concept of Irenimus Pascoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30 Chalepistes Brown, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Abstract The taxonomy of the New Zealand weevil genus Irenimus Pascoe, 1876 is revised, resulting in a narrower concept of the genus than has been considered in recent decades. In total, the genus now contains only seven species. In addition to the type species, I. parilis Pascoe, 1876, the genus contains I. duplex (Broun, 1904) and five newly described species: I. an- iptus new species (type locality, Oamaru, DN), I. crinitus new species (type locality, Hakataramea Valley, SC), I. minimus new species (type locality, Alexandra, CO), I. stichus new species (type locality, Tekapo, MK) and I. thoracicus new spe- cies (type locality, Oamaru, DN). The genus Chalepistes new genus is established to contain the majority of species pre- viously described in the genus Catoptes Schönherr, 1842, but also including species described in Brachyolus White, 1846; Irenimus Pascoe, 1876; Inophloeus Pascoe, 1875; and Nicaeana Pascoe, 1877. A total of 27 valid described species are new combinations with Chalepistes: C. aequalis (Broun, 1895) (from Irenimus), C. albosparsus (Broun, 1917) (from Ire- nimus), C. apicalis (Broun, 1923) (from Catoptes), C. asperatus (Broun, 1914) (from Brachyolus), C. compressus (Broun, 1880) (from Irenimus), C. costifer (Broun, 1886) (from Inophloeus), C. curvus (Barratt & Kuschel, 1996) (from Irenimus), C. dehiscens (Broun, 1917) (from Catoptes), C. dugdalei (Barratt & Kuschel, 1996) (from Irenimus), C. egens (Broun, 1904) (from Irenimus), C. inaequalis (Sharp, 1886) (from Brachyolus), C. instabilis (Marshall, 1931) (from Catoptes), C. latipennis (Broun, 1893) (from Catoptes), C. limbatus (Broun, 1909) (from Catoptes), C. lobatus (Broun, 1921) (from Catoptes), C. patricki (Barratt & Kuschel, 1996) (from Irenimus), C. pensus (Broun, 1914) (from Inophloeus), C. placidus (Broun, 1914) (from Nicaeana), C. posticalis (Broun, 1893) (from Irenimus), C. rhesus (Pascoe, 1875) (from Inophloeus), C. rubidus (Broun, 1881) (from Inophloeus), C. similis (Barratt & Kuschel, 1996) (from Irenimus), C. spectabilis (Broun, 1914) (from Catoptes), C. spermophilus (Broun, 1895), revised status (from Irenimus), C. stolidus (Broun, 1886) (from Accepted by R. Anderson: 17 Feb. 2017; published: 8 May 2017 1 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 Irenimus), C. tenebricus (Broun, 1893) (from Catoptes), C. vastator (Broun, 1893) (from Irenimus). Numerous new syn- onyms with species of Chalepistes are also proposed: Brachyolus fuscipictus Broun, 1914 and Brachyolus terricola Broun, 1917 are junior subjective synonyms of Chalepistes asperatus (Broun); Brachyolus cervalis Broun, 1903 and Brachyolus sylvaticus Broun, 1910 are junior subjective synonyms of Chalepistes costifer (Broun); Inophloeus tricostatus Broun, 1915 is a junior subjective synonym of Chalepistes pensus (Broun); Catoptes pallidipes Broun, 1917, Catoptes flaviventris Broun, 1917 and Catoptes nigricans Broun, 1917 are junior subjective synonyms of Chalepistes placidus (Broun); Inophloeus longicornis Broun, 1904, Inophloeus medius Broun, 1893, Inophloeus sulcicollis Broun, 1914 and Inophloeus suturalis Broun, 1893 are junior subjective synonyms of Chalepistes rhesus (Pascoe); Inophloeus albonotata Broun, 1893, Catoptes asperellus Broun, 1893, Irenimus bicostatus Broun, 1886, Catoptes caliginosus Broun, 1893, Catoptes chalmeri Broun, 1893, Catoptes decorus Broun, 1893, Inophloeus discrepans Broun, 1904, Catoptes fumosus Broun, 1914, Catoptes furvus Broun, 1893, Catoptes humeralis Broun, 1893, Catoptes longulus Sharp, 1886, Inophloeus nigellus Broun, 1881, Irenimus pilosellus Broun, 1886 and Catoptes scutellaris Sharp, 1886 are junior subjective syn- onyms of Chalepistes rubidus (Broun); Catoptes subnitidus Broun, 1914 and Catoptes curvatus Broun, 1914 are junior subjective synonyms of Chalepistes spermophilus (Broun); Catoptes brevicornis Sharp, 1886 and Catoptes vexator Broun, 1904 are junior subjective synonyms of Chalepistes stolidus (Broun); and Catoptes aemulator Broun, 1893 and Catoptes argentalis Broun, 1914 are junior subjective synonyms of Chalepistes tenebricus (Broun). Additional new com- binations include Inophloeus robustus (Broun, 1917) (from Catoptes) and Nicaeana fraudator (Marshall, 1931) (from Catoptes), while Catoptes postrectus Marshall, 1931 is a new synonym of Protolobus obscurus Sharp, 1886. Introduction Anthropogenic modification of habitats usually results in decreased species richness, particularly of indigenous elements of the system (Harris & Burns, 2000; Murphy & Romanuk, 2014). The lowland environments in New Zealand have undergone dramatic changes since human settlement, especially in the 200 years since European colonisation (Ewers et al., 2006). Much of the New Zealand lowlands has been converted into agricultural pasture based on a simplified flora of exotic fodder crops (Leathwick et al., 2003; Goldson et al., 2014). The persistence of endemic New Zealand faunal elements in these highly modified environments is therefore of interest. Broad-nosed weevils in the genus Irenimus Pascoe, 1876 and related genera are examples of New Zealand endemics that have survived in agricultural environments. As a result, they have been the subject of much research into their ecology, due to some species achieving minor pest status (Baker, 1980; Scott, 1984; Stewart, 1981) and because of concern over the effects of non-target parasitism by braconid wasps introduced to control adventive pest weevils (Barratt, 2004; Barratt et al., 2000). The taxonomy and systematics of the group has, however, received little attention since the pioneering work of Thomas Broun, resulting in confusion regarding nomenclature and the limits of species and genera (Bremner, 1988; Barratt & Kuschel, 1996; Macfarlane et al., 2011). Irenimus was described by Pascoe (1876) as a monotypic genus containing I. parilis Pascoe, 1876. Subsequently, Broun (1886) described four species in Irenimus, before Sharp (1886) synonymised the genus with Catoptes Schönherr, 1842. This synonymy was accepted by Broun (1910). Kuschel (1969), having discovered that Catoptes had been misapplied by Broun, reinstated Irenimus Pascoe and placed Catoptes aequalis Broun, 1895 and Catoptes compressus Broun, 1880 into Irenimus. He also recommended that, until a comprehensive revision could be undertaken, all species described by Broun, Sharp and Marshall under Catoptes should provisionally be placed in Irenimus. Kuschel’s (1969) concept of Catoptes Schönherr, with generic synonyms of Asaphia Broun, 1881, Empaeotes Pascoe, 1876, Exonastus Broun, 1921, Getopsephus Broun, 1913, Platyomida White, 1846, Protophormus Sharp, 1886 and Tigones Broun, 1886 is followed in this paper. With the restricted definition of Irenimus presented here, the new genus Chalepistes is made available to accommodate a number of species that would otherwise be unplaced to genus. A monograph of Chalepistes is in preparation, which will include keys to species, type details and descriptions of new species. In the interim, a list of species placed in Chalepistes is provided, with synonyms. 2 · Zootaxa 4263 (1) © 2017 Magnolia Press BROWN Methods Field-collected specimens were killed in 100% ethanol or placed directly into a freezer at a temperature of -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 cleared by digestion for c. 36 hours in a porcine pancreatin solution (Álvarez Padilla & Hormiga, 2008). The lysate from this digest was then used for DNA extraction. If specimens were unsuitable for DNA extraction, or had not cleared satisfactorily at the end of this time, abdomens were digested in cold 10% KOH for up to two hours. After clearing, the abdomen was flayed by cutting down through the right margin of the membranous tergites 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 1% w/v solution of Chlorazol Black dissolved 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) (Liberti, 2005), which was then pinned below the specimen. Terminology follows Oberprieler et al. (2014), Lawrence et al. (2010) and Arzanov (2003). Body length was measured in lateral view, from the anterior margin of the eyes to the elytral declivity. Rostrum width was measured across the antennal insertions. Legs are described in their idealised laterally extended position, thereby having dorsal, ventral, anterior and posterior surfaces. The term ‘pappolepida’ is coined (after the Greek pappos, ‘downy’, Brown 1956) to describe a distinctive type of appressed scale found in abundance on the venter of several species. Pappolepida are multiply divided, giving them a feathery or velvety appearance (Figure 38). Descriptions of colour follow the terminology provided by the National Bureau of Standards (Kelly & Judd, 1976). The NBS centroid colours are a dictionary of 267 colours, with natural-language descriptions. Digital representations of these colours have been provided by Jaffer (2011). Specimens were loaned from and deposited in the following collections: AMNZ Auckland War Memorial Museum, Auckland, New Zealand ANIC Australian National Insect Collection, CSIRO, Canberra, Australia CMNZ Canterbury Museum, Christchurch, New Zealand BMNH Natural History Museum, London, United Kingdom FRNZ National Forest Insect Collection, Scion, Christchurch, New Zealand IACC Invermay Agricultural Centre Collection, Mosgiel, New Zealand LUNZ Lincoln University Entomology Research Collection, Lincoln, Canterbury, New Zealand MONZ Te Papa Tongarewa, National Museum of New Zealand, Wellington, New Zealand NZAC New Zealand Arthropod Collection, Manaaki Whenua Landcare Research, Tamaki, Auckland, New Zealand OMNZ Otago Museum, Dunedin, New Zealand USNM Smithsonian Institution National Museum of Natural History, Washington D.C., United States of America DNA sequencing and analysis. Only freshly collected specimens were used for sequencing. Genomic DNA was extracted from 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 (Table 1). 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 MgCl and 0.2 μM of forward and reverse 2 primers. The COI primer combination LCO1490-JJ/TL2-N-3014 was used preferentially, in order to amplify the whole gene, which was then sequenced using all four primers. If amplification using this combination was not successful, C1-J-2183/TL2-N-3014 was used. Reactions were run on a C1000 Touch thermal cycler (Bio-Rad REVISION OF IRENIMUS WEEVILS Zootaxa 4263 (1) © 2017 Magnolia Press · 3 Laboratories Inc., Hercules, CA, USA) or a MJ Mini thermal cycler (Bio-Rad Laboratories Inc.) with an initial denature of 94°C for 2 min, followed by 40 cycles of denaturing 94°C (20 s), variable annealing temperature (20 s) and extension 72°C (60 s), with a final extension at 72°C for 5 min. Annealing temperatures were 45°C for COI and 52°C for 28S reactions. ArgK and CAD reactions were amplified using a touchdown protocol, with annealing temperatures starting at 50°C, decreasing by 1°C per cycle for 5 cycles followed by 35 cycles at 45°C. Purified PCR products were sequenced by Macrogen (Seoul, Korea) using ABI BigDye 3.1 technology on an ABI3730XL platform (Applied Biosystems). TABLE 1. Markers and PCR primer combinations used in this research. Marker Primer name Direction Primer sequence Reference COI C1-J-2183 Forward 5′-CAA CAT TTA TTT TGA TTT TTT GG-3′ Simon et al. 1994 LCO1490-JJ Forward 5′-CHA CWA AYC ATA AAG ATA TYG G-3′ Astrin & Stüben 2008 HCO2198-JJ Reverse 5′-AWA CTT CVG GRT GVC CAA ARA ATC A-3′ Astrin & Stüben 2008 TL2-N-3014 Reverse 5′-TCC AAT GCA CTA ATC TGC CAT ATT A-3′ Simon et al. 1994 28S S3660 Forward 5′-GAG AGT TMA ASA GTA CGT GAA AC-3′ Sequeira et al. 2000 28S-Ff Reverse 5′-TTA CAC ACT CCT TAG CGG AT-3′ Gómez-Zurita et al. 2005 ArgK ArgKforB4 Forward 5′-GAY CCC ATC ATC GAR GAC TAC C-3′ McKenna et al. 2009 ArgKrevB1 Reverse 5′-TCN GTR AGR CCC ATW CGT CTC-3′ McKenna et al. 2009 CAD CADfor4 Forward 5′-TGG AAR GAR GTB GAR TAC GAR GTG GTY Jordal et al. 2011 CG-3′ CADrev1mod Reverse 5′-GCC ATY RCY TCB CCY ACR CTY TTC AT-3′ Jordal et al. 2011 Sequences were aligned by eye in SEAVIEW (Guoy et al., 2010), trimmed to minimise missing data, and analysed in R (R Development Core Team, 2010). Uncorrected genetic distances (p-distances) were calculated using APE (Paradis et al., 2004), which were then decomposed into interspecific and intraspecific components using SPIDER (Brown et al., 2012). Diagnostic nucleotides (Sarkar et al., 2008) for each species were identified using SPIDER. Illustrations. Line drawings were made with a Zeiss Stemi SV6 stereo microscope fitted with a camera lucida. The drawings were scanned and inked digitally and saved in vector graphic format using INKSCAPE (v. 0.91, Inkscape Team, 2004-2017). Genitalia illustrations were prepared from photographs. Habitus photographs were created using focus stacking, as implemented in the software Nikon NIS ELEMENTS (v. 4.10). The image stack was prepared using a Nikon AZ100M stereomicroscope fitted with a Nikon DS-Ri 1 digital camera and mechanical z-stepper. The top and bottom planes of the specimen was specified in NIS ELEMENTS, which then took multiple photographs and combined them into a single image with a large depth of field. Specimen data. 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 is given in square brackets ([…]). Two-letter regional codes (DN, SC, CO, etc) follow those proposed by Crosby et al. (1998). Coordinates given after the locality names are in the WGS84 datum. Coordinates tagged “R” (Recorded) were obtained from coordinates on the label when given, or from consultation with the collector. Coordinates tagged “A” (Approximate) were determined by using available gazetteers, primarily the New Zealand Gazetteer of Place Names (Land Information New Zealand, 2016). These georeferenced data were used to extract estimated altitudes from 25 m resolution digital elevation models of New Zealand provided by Landcare Research (Landcare Research, 2010a, 2010b). Species and taxonomic concepts. In common with usage throughout history, a generative conception of species is used here (Wilkins, 2009). In this paper, I infer the extent of a species by the formation of diagnosable entities as revealed by morphological traits. Genetic data provide extra evidence that the species described herein represent separate lineages. These taxa are proposed in order to investigate the evolutionary processes leading to their existence and the ecological functions that they fulfil (Wilkins, 2010). 4 · Zootaxa 4263 (1) © 2017 Magnolia Press BROWN Alignments between changing historical taxonomic concepts of species and genera were created using the EULER/X toolkit (Chen et al., 2014; Jansen & Franz, 2015; Franz et al., 2015). This software evaluates logical consistency between two taxonomies, given a list of the region connection calculus (RCC-5) relations (Li & Ying, 2003; Franz et al., 2015) between the two. Irenimus Pascoe, 1876 Irenimus Pascoe, 1876:54. Synonymised with Catoptes Schönherr by Sharp, 1886:422. Reinstated by Kuschel, 1969:799. Type species: Irenimus parilis Pascoe, 1876:54, by monotypy. Gender: masculine. Diagnosis. Integument densely covered with round or oval appressed scales. Rostrum short and stout, around 1.5 times longer than wide; subparallel proximally. Epistome indistinctly differentiated from frons. Scrobes lateral. Ventral curvature of head angulate. Elytra without tubercles or protuberances. Metanepisternum narrow, suture complete. Metafemora without sharp demarcation between scaled and unscaled regions on posterior surface. Metatibiae with a narrow to wide corbel and a distinctively shaped, angulate apical comb (Figures 29–33). Penis with a sclerotised dorsal membrane. Differential diagnosis. Species of Irenimus can be distinguished from other New Zealand weevils by the combination of characters given above. The indistinct epistome and angulate ventral head curvature distinguishes them from Catoptes, which have a distinctly demarked epistome and a rounded ventral head curvature, with the angle between head and rostrum approximately 140°. The complete metanepisternal suture and the form of the metatibial apex distinguish Irenimus from Chalepistes, which has the metanepisternal suture invisible, a simple metatibial apex, and a sharply demarked bare area on the posterior surface of the metafemora. The convex elytral disc and lack of elytral tubercles or prominences distringuish Irenimus from Inophloeus villaris Pascoe and allied species which have a flat elytral disc, formed by a subcarinate interstriae 7, and strong protuberances on interstriae 5 above the declivity. The wide, readily visible metanepisternum and the discrete appressed scales on the elytra distinguish Irenimus from Protolobus, which has a very narrow metanepisternum that is often obscured by the lateral margin of the elytra and has dull, strongly overlapping, inseparable appressed scales on the elytra. The proximally subparallel rostrum and laterally situated scrobes distinguish Irenimus from Haplolobus and Nicaeana, both of which have proximally expanding rostra and dorsally situated scrobes. Description. Body length ranging from 3.2 mm to 6.9 mm. Densely covered with scales on all surfaces. Rostrum. Subparallel proximally, abruptly widened at antennal insertions. Epistome plurisetose, slightly raised above frons but indistinctly differentiated. Epifrons with longitudinal medial carina, lacking sulci; continuous with occiput, without distinct dorsal separation between head and rostrum. Antennae. Sockets dorsolateral, sited in apical 1/3 of rostrum. Scape clavate, reaching to or exceeding the middle of the eye. Funicular segments clavate, subspherical or oblately spheroid, moderately to loosely articulated, segment 7 almost as wide as club. Club two times longer than wide, tapering apically. Head. Width of vertex between eyes greater than width of rostrum. Eyes large, lateral, flat, ovate to subcircular with long axis vertical, parallel with sagittal axis. Ventral curvature of head and rostrum angulate, approximately 90°. Pronotum. Disc smooth, evenly convex. Postocular lobes poorly to well developed, vibrissae present. Elytra. Approximately parallel-sided in anterior 2/3. Elongate scales arising from interstriae. Elytral declivity rounded. No interstriae formed into tubercles, at most a slight swelling on interstria 3 above the elytral declivity. Ventral margin sinuous, highest point around level of metacoxae. Thorax. Procoxae contiguous. Prosternum visible behind procoxae as a raised tubercle (“prosternellum”). Metaventrite with medial suture visible only as a small, circular fovea posteriorly. Abdomen. Ventrites 1 and 2 fused, ventrites 3 to 5 free. Ventrite 1 and 2 subequal in length at midpoint; ventrites 3 and 4 subequal in length, approximately 0.5 times length of ventrites 1 and 2; ventrite 5 approximately equal in length to ventrites 1 and 2. Suture separating ventrites 1 and 2 curved anteriorly at midpoint, 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 depth around distal 1/4. Tibiae with indistinct denticles along ventral margin of protibiae and mesotibiae; mucrones present on protibiae and mesotibiae; protibiae wider in distal 1/3 than proximal 1/3, incurved at apex; metatibiae with subparallel dorsoventral margins; apical comb angulate with pale setae; corbel present, narrow to wide, bare. Tarsi with long, coarse setae on dorsal surface, without appressed scales; underside with short, dense setae forming pads. Claws simple, separate, diverging. Male genitalia. Hemisternite 8 divided, with a paired membranous lobe on the anterior margin of the membrane joining the two halves. Penis tubular, strongly curved; lateral lobes REVISION OF IRENIMUS WEEVILS Zootaxa 4263 (1) © 2017 Magnolia Press · 5 FIGURES 1–8. Habitus photographs of Irenimus males. 1, 2: I. aniptus. 3, 4: I. crinitus. 5, 6: I. stichus, holotype. 7, 8: I. minimus. Scale bars = 1 mm. 6 · Zootaxa 4263 (1) © 2017 Magnolia Press BROWN FIGURES 9–14. Habitus photographs of Irenimus males. 9, 10: I. duplex. 11, 12: I. parilis. 13, 14: I. thoracicus, holotype. Scale bars = 1 mm. separated dorsally by sclerotised dorsal membrane extending from ostium to anterior margin (Figure 41, dm); temones longer than penis. Endophallus moderate in length, usually reaching anterior 1/3 of temones when in repose; armed with large aggonoporial sclerite, other sclerites variably present. Tegmen with ring complete; parameroid lobes well-developed; manubrium shorter than temones. Female genitalia. Sternite 8 with spiculum ventrale over 2.5 times as long as blade. Gonocoxite divided into two parts; proximal gonocoxite around 2.6 times longer than distal gonocoxite, largely unsclerotised except for strongly sclerotised rod; distal gonocoxite lightly sclerotised. Bursal sclerites present or absent, numbering 2 if present. Distribution. All species found in Canterbury and Otago, South Island, New Zealand, with a disjunct population of I. parilis in the central North Island. Etymology. Pascoe (1876) did not explain how the name was derived. It may have come from the Greek eirene, meaning ‘peace’, and the Latin suffix -imus meaning ‘having the quality of’. Remarks. The extensive dorsally sclerotised membrane of the penis is a strong apomorphy for Irenimus. However, all external characters that distinguish the genus are considered plesiomorphic. REVISION OF IRENIMUS WEEVILS Zootaxa 4263 (1) © 2017 Magnolia Press · 7 FIGURES 15–22. Habitus photographs of Irenimus females. 15, 16: I. aniptus. 17, 18: I. crinitus. 19, 20: I. stichus. 21, 22: I. minimus. Scale bars = 1 mm. Key to species of Irenimus 1 Scape clothed with setae only. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 - Scape clothed with appressed scales as well as setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 2 (1) Ventral margin of metatibial apex with a strong, conspicuous emargination. Elytra with elongate scales mostly decumbent. Abdominal ventrite 1 with few pappolepida, clothed primarily with appressed scales . . . . . . . . . . . . . . . . . . . .. I. stichus n. sp. - Ventral margin of metatibial apex entire or with an inconspicuous emargination. Elytra with elongate scales semi-erect to erect. Vestiture of abdominal ventrite 1 dominated by numerous pappolepida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 (2) Elongate scales of pronotum of equal length. Dorsal appressed scales with coarse ridges, resulting in a dull appearance. 8 · Zootaxa 4263 (1) © 2017 Magnolia Press BROWN Metaventrite with appressed scales laterally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. aniptus n. sp. - Elongate scales of pronotum longer towards anterior margin. Dorsal appressed scales with fine ridges, resulting in a glossy appearance. Metaventrite with pappolepida laterally (Figure 38). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. crinitus n. sp. 4 (1) Larger species, body length longer than 4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 - Smaller species, body length shorter than 4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. minimus n. sp. 5 (4) Metatibiae with broad corbel (Figure 31). Ventral margin of metatibial apex emarginate (Figure 31). Pronotum with appressed scales flat, resulting in a smooth appearance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. parilis Pascoe - Metatibiae with narrow corbel (Figure 33). Ventral margin of metatibial apex entire (Figure 33). Pronotum with appressed scales domed, resulting in a bumpy appearance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6 (5) Lateral margins of pronotum evenly curved along length (Figure 39) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. duplex (Broun) - Lateral margins of pronotum strongly curved in anterior 2/3; subparallel in posterior 1/3 (Figure 40) . . . . I. thoracicus n. sp. FIGURES 23–28. Habitus photographs of Irenimus females. 23, 24: I. duplex. 25, 26: I. parilis. 27, 28: I. thoracicus. Scale bars = 1 mm. REVISION OF IRENIMUS WEEVILS Zootaxa 4263 (1) © 2017 Magnolia Press · 9 FIGURES 29–34. Left metatibia, anterior view. 29, Irenimus minimus; 30, Irenimus parilis; 31, Irenimus parilis, apical view; 32, Irenimus duplex; 33, Irenimus duplex, apical view; 34, Chalepistes compressus. Abbreviations: cl–corbel; sc—setal comb; ve—ventral emargination. Drawings not to scale. FIGURES 35–36. Pronotum, left lateral view. 35, Irenimus crinitus; 36, Irenimus stichus. Abbreviations: atg—anterior transverse groove; ptg—posterior transverse groove; tooth—pronotal tooth. 1. Irenimus aniptus Brown, new species Figures 1, 2, 15, 16, 41, 42, 43, 44, 45, 46, 85 Diagnosis. Medium-sized species; oval; grey, often with encrustation of soil. Scape without appressed scales. Metaventrite laterally with appressed scales. Elytra with inconspicuous swelling on interstriae 3. Appressed scales with coarse ridges, resulting in a duller appearance than I. crinitus n. sp. 10 · Zootaxa 4263 (1) © 2017 Magnolia Press BROWN