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SE IES E I O AU A O EW EA A M C u a (d d n 199 a ca e esea c Mou A e esea c Ce e i a e ag 2 0, Auck a , ew ea a Fauna of New Zealand Ko te Aitanga Pepeke o Aotearoa
Number / Nama 39
Molytini (Insecta: Coleoptera: Curculionidae: Molytinae)
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Otago Museum P.O. Box 6202, Dunedin, New Zealand
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Class Insecta Order Coleoptera Family Curculionidae Subfamily Molytinae Tribe Molytini
Molytine weevils I us a io / W akaa ua a am us u e cu a us (Il- l tr t r / K - h h l r These flightless weevils are some of the largest beetles in New Zealand, and they are some of the largest members in ā ī h r M l t n the world of their tribe Molytini. Adult speargrass weevils Κ ēn ī h r r r - r ēt h n āā pīt r r h r t (genus y e o ius can range in colour from a red-brown A t r r t rāt nō ēt h n t n to black, and many species have elegant white stripes on r h tō rāt ī n M l t n Ιn t r h n ā their backs, whereas adult karo and knobbled weevils ī h r n h t r ( t p n n Lyperobius), n ā (genus a am us are predominately dark brown with t n ā p h ō ō h p n n triangular or rounded tubercles on their sides and backs. rān t Ot rā rā nō ēt h h āt h t This study redescribes the twelve known species and tā h r t ā t p r r ō rāt t rā K provides descriptions of eight new speargrass weevils. n ā p t ī h r n h t h t r t ī h r How to identify these weevils is covered in detail, and pūr r (nō t p n n Hadramphus), he parauri te tae, there is a key to the known and new kinds. Habitus t rā t t rā n ā ēt h t n h p r h t h drawings of these weevils, as well as detailed illustrations t p t r rān t āh of key external and internal features, are provided. I r t tēn r n h t n h h t ā- p As the popular name speargrass weevil implies, adults of n ā ī h r M l t n t ā ōh t n most species feed on the rigid spiky leaves and large t t ēt h h r K t ēn h flowering heads of speargrasses or spaniards (taramea), h ī h r n h t r K h t nō n ā ōr r and they are sometimes also found on kopoti. Grubs of n ā h ār ōh t t t h tēnā most species feed on roots and the growing leaf crown of tēnā Arā nh ēt h h h ēn ī h r these members of the carrot family (Apiaceae), whereas ēt h h h n ā āh t n t ō rāt t n n the karo weevil grubs bore in the trunks of karo trees on the h r t nō h Poor Knights Islands. K t n ār n r n t n ēn t h n These weevils are known to occur in insular habitats from t t r t t n t n n n p tēn the subtropical Poor Knights Islands off the Northland ī h r ; n n ā r t r t r n ā p coast, to the isolated Chatham Islands in the east, and the t t r nō t nō ēt h n h t h n cold subantarctic Snares Islands in the south. They are t t p ī n t p t K n ā t r n ū most numerous and diverse in the South Island mountains, t n n n ā ī h r t r n ā with maximum species diversity in West Otago and n ā r h ēn h ān t hān ār t ( t Fiordland. Ap t n pūt En r nō n ā t r n ū t An attempt to reconstruct the genealogy of these weevils ī h r n h t h t r — tā rāt h h r is presented. Using this genealogy as a basis, the h r h h r n ā h n ā r h t ecological and biogeographic history of their divergence h
(co i ue o e ea (a a ae e o u
-5- in New Zealand is estimated. It is suggested that they are Kitea ēn ī h r M l t n ēt h ā n t r t h an old group, of at least 65 million years duration in n ā pārūrū t r r r n n ā Aotearoa, descended from a widespread ancestral t r h t h Wh r r n h tār complex of coastal, insular, lowland, and mountainous n t ā h t p t t n ā āh r r pērā sites that have long been conspicuous features of our ā M t r Sn r t t n Kā r h āh t t country. n ā n W p n ō t h h n ā ī h r These weevils are generally considered in New Zealand t t t h h n ā ī h r Ā t āh conservation biology as threatened species that are highly r r t t n h h t n ēn pīt r t t ā vulnerable to the combined impact of rodent predators and Ōtā t Ur t Kār h t habitat modification. One species appears to have become K h t t h h t t r t h p p extinct since European settlement. Insect ecologists who n ā ī h r n ā h h t h p p n have made detailed studies of the knobbled weevil h tūāp p h t h t h h h r ēn consider that they are good indicators of whether there are r t n ā t K t r h t āh tā rāt n h t h t rats on an island. t ēt h t n ā t n ān t hōr p h r n ā ī h r t t t h n r t n ā t n t n E h p t n nō t r hā n ā ī h r Contributor r b n Cr is Honorary Curator of n — rā n t t 5 r n t n h n Entomology at the Otago Museum in Dunedin. His past A t r E h p t n nō h n h ō rāt research has included systematic revisions of tussock tīp n n ā āh h t n — t t h n tēnā butterflies and geometrid moths, as well as many n ā t r ē ā t h tār tēnā n ā h n publications on the history, method, and theory of t p t p t tēnā t t n ā p n I r t n ā h r hī n ā t n ān A t r biogeography and phylogenetic systematics. Currently he h t n t n n h pāh h t āh t is working on collecting and describing new species of n h ēn pīt r t E n h ōr r boulder, copper, and black mountain butterflies, and the n h t n n ā āh t n n rāt conservation status of katipo spiders. pēn t r tō rāt t n h r r n ā h Ko te t h n ā t b n Cr t K t t h r r ē t āh ō rāt ā n n h Arā tēt h Wāh n Māt n ā A t n p t Wh r p r h r t n n n n r t t n n Ōtā Ōt p t K ēt h n r n h t n t ā hā ērā ēt h t h n t r t r n ā t n n ā t r t t r t r nō t h h n ā p p āt r n h ί t ī h r pūr r ā h rōpūt n n ā pūr r h pātītī t r n n ā tā rāt n h tēn ī h r h t h pūr r h ` tr d K h p t t h n h pā h n h n tēt h t r t r ā r ana ki te hīt r n ā t n n ā ātāp n t r - rān (Arā n h t n n ā ī h r pūr r t h n n ā pūn h h rōpū pā n t r p h r ā h r r n h ī h r kunenga mai ēt h K tān h ēn rā h h r r p t h t r he whakaahua i ēt h pūr r h h K ēn hou, nō r t n ā hān pūr r h n h t n ā pūr r h r n ā pūr r h n h n h pān te t Arā nō tēt h p p hā n t t r t r i te ongeonge haere o te t pō
K p Ā h n - Gl r aitanga a Pepeke — generic term for insects tā — tr p d koiora-matawhenua — biogeography tār — l t d r t ō ō — r d-br n Tawhiti Rahi — Poor Knights Islands kunenga — evolution Te Taka o te Kār h t — rdl nd monemone noa — obliterated, rendered extinct tona — tubercle ōr r — xp d t d n r t r hā — v r n nt t ongeonge — becoming rare, endangered whenua tapotupotu — lowland pāh h — n rt n n t bl ī h r — v l pātītī t r n — t r Translation by b pūr r — n bbl d Huatau Consultants, Wellington
—6— A S AC New Zealand's coxella, karo, knobbled, and speargrass weevils are revised. Sixteen species are recognised in the genus Lyperobius Pascoe, 1876, of which eight are new (australis, barbarae, clarkei, eylesi, glacialis, montanus, patricki, and townsendi). Four species are recognised in the genus Hadramphus Broun, 1911. Karocolens Kuschel, 1987, based on K. pittospori Kuschel, 1987, is reduced to synonymy with Hadramphus, and the new combination H. pittospori is proposed. Lyperobius aciphyllae Broun, 1917 is a subjective synonym of L. cupiendus Broun, 1886. Genera and species are described, and a key to adults is given. Life history, biology, and conservation status and value are reviewed. A numerical phylogenetic analysis exploring the relationships of the two genera and their included species is shown to contain significant phylogenetic structure, against a null hypothesis of random noise, by bootstrapping and tree-length distribution skewness. Biogeographical and ecological history, including host plant shifts, are discussed.
Craw, R.C. 1999. Molytini (Insecta: Coleoptera: Curculionidae: Molytinae). Fauna o New Zealand 39, 68 pp. C ECK IS O A A Phylogenetic analysis 11 Methods 11 20 Genus Hadramphus Broun, 1911 Monophyly of in-group clade Hadramphus + Karocolens Kuschel, 1987, new synonymy Lyperobius 11 pittospori (Kuschel, 1987), new combination 21 Character list and discussion of character-state spinipennis Broun, 1911 21 polarity 11 stilbocarpae Kuschel, 1971 22 Phylogenetic results and relationships 13 tuberculatus (Pascoe, 1877) 22 Statistical tests of phylogenetic structure 15 Genus Lyperobius Pascoe, 1876 23 Historical ecology and biogeography 16 australis new species 23 Methods 16 barbarae new species 24 Plant-host analysis 16 carinatus Broun, 1881 24 Vicariant distribution and sympatric occurrences 17 clarkei new species 25 Elevational range, habitat, and biogeographic coxalis Kuschel, 1987 26 history 17 cupiendus Broun, 1886 26 Conservation status and value 19 aciphyllae Broun, 1917 new synonymy Descriptions (see `Checklist of Taxa') 20 eylesi new species 27 References 32 fallax Broun, 1917 27 Appendix 1: Offshore Molytini examined 37 glacialis new species 28 Key to genera and species of Molytini known from hudsoni Broun, 1914 28 New Zealand 37 huttoni Pascoe, 1876 29 Illustrations 39 montanus new species 30 Distribution maps 57 nesidiotes Kuschel, 1987 30 Taxonomic index 62 patricki new species 31 Appendix 2: Host-plant associations 63 spedenii Broun, 1917 31 townsendi new species 32 ACK OW E GME S For the loan of specimens I thank Mr J.W. Early and Mr CO E S K.A.J. Wise (AMNZ), Dr R.M. Emberson (LUNZ), Mr Introduction 8 A.C. Harris (OMNZ), Dr C.H.C. Lyal (BMNH), Mr R.L. Taxonomic definition of N.Z. Molytini 8 Palma (MONZ), and Mr A. Savill (formerly of CMNZ). Morphological characters 9 Valuable material of offshore Molytini was made avail- Life history and biology 10 able by Dr R.S. Anderson (Canadian Museum of Nature),
-7- Dr K. Morimoto (Kyushu Univerity, Fukuoka, Japan), and Although they are popularly and frequently known as Dr C.W. O'Brien (Tallahassee, Florida, U.S.A.). `giant weevils', this term was originally coined (in a New Particular thanks are due to Dr B.I.P. Barratt (BIPB) Zealand context) as a collective common name for the (AgResearch, Invermay Agricultural Centre, Mosgiel) large subantarctic species of adelognathous (i.e., broad- and Mr B.H. Patrick (BHPC) (formerly of Department of nosed) weevils at present placed in the genera Heterexis Conservation, Dunedin, now OMNZ) for advice, Broun and Oclandius Blanchard (Hudson 1934). Spear- encouragement, information, and loan of specimens grass weevil (for all species of Lyperobius), coxella weevil without which this revision would never have been (for Hadramphus spinipennis), karo weevil (for H. pitto- completed. Dr R. Gray (University of Auckland) gave spori), and knobbled weevil (for H. stilbocarpae) are more extensive advice and assistance concerning phylogenetic accurate common names, and have been used in recent analysis. Correspondence with Dr N. Platnick (American popular treatments (e.g., Meads 1990, Foord 1992). Museum of Natural History, New York) helped clarify Previous systematic studies of New Zealand Molytini nomenclatural implications of the phylogenetic analysis. have been isolated descriptions of genera and species Mr B.W. Thomas (Landcare Research, Nelson) provided (Pascoe 1876, 1877; Broun 1881, 1886, 1911, 1914, new information on Hadramphus. Referees Dr J.J. 1917; Kuschel 1971, 1987). Larvae and pupae of some Morrone (Universidad Nacional de La Plata, Argentina) species have been described (May 1971, 1981, 1987a, and Mr I. Millar (Nelson/Marlborough Conservancy, 1993). Bull (1967) provided a detailed study of the Department of Conservation, Nelson) made useful morphology, life history, and biology of L. huttoni. The comments on the manuscript. Special thanks to Mr D.W. present study is based on examination of more than 1300 Helmore (Landcare Research, Auckland) for his fine adult specimens. habitus studies and drawings of external morphology. Abbreviations used above and in citations of speci- mens examined: A O OMIC E I I IO O MO Y I I AMNZ Auckland Institute and War Memorial Museum, Weevils belonging to the tribe Molytini of the curculionid Auckland, N.Z. subfamily Molytinaecan be recognised in the New Zea- BHPC Brian Patrick private collection, Dunedin, N.Z. land fauna, and distinguished from offshore Molytini by [now in OMNZ] the following combination of characters. BIPB Dr Barbara Barratt private collection, Dunedin, N.Z. Adults. Length (excluding rostrum) 11.5-28 mm, greatest BMNH British Museum (Natural History), now The width (across elytra) 5.9-12.5 mm. Large, brachypterous, Natural History Museum, London, England flightless molytine curculionids. Wings rudimentary, nar- CMNZ Canterbury Museum, Christchurch, New Zealand rowly laminate, 0.2-0.65x as long as elytra. Antennae LUNZ Dept of Entomology, Lincoln University, Lin- geniculate; funicle with 7 segments, the 7th free from club; coln, Canterbury, N.Z. club with 3 segments, the lst longer than remainder of MONZ Museum of New Zealand, Wellington, N.Z. club, the 3rd with a transverse line; sutures straight, ob- NZAC New Zealand Arthropod Collection, Landcare lique. Prothorax with ocular lobes prominent to obsolete, Research, Mt Albert Research Centre, Auck- prosternum emarginate. Procoxae contiguous, metacoxae land, N.Z. transverse. Tibial apex with uncus and premucro, the OMNZ Otago Museum, Dunedin, N.Z. uncus arising from a low corbel carina (Fig. 35); distal protibial comb rounded, meso- and metatibial combs ob- liquely sinuate. Tarsi spongy beneath; 1st segment elon- gate; 3rd segment much broader than the 2nd, bilobed; 4th I O UC IO segment concealed; 5th segment longer than the 3rd; claws New Zealand weevils assigned to the tribe Molytini (= symmetric, free. Liparini) comprise some of our largest beetles, and some Male. Tergite 8 not concealed by tergite 7. Sternite 8 species are among the largest in the tribe. They are con- composed of 2 separate sclerotised plates, posteriorly spicuous members of insular, montane, subalpine, and emarginate, with short latero-apical setae. Sternite 9 with alpine communities. Twenty species in two genera — asymmetric arms and a sclerotised lobe (Fig. 44, 46). Teg- Lyperobius Pascoe, 1876 and Hadramphus Broun, 1911 — men complete; parameres well developed, with a short are recognised in this study, but further undescribed manubrium (Fig. 45, 47). Aedeagus: median lobe weakly species may occur, particularly in the northern half of the sclerotised along dorsal median line (Fig. 58); pedal por- South Island. tion divided laterally by a greatly distensible membrane —8— (Fig. 64). Endophallus non-eversible, attached to apical Hudson (1934) and Kuschel (1987) this tribe, as margin of aedeagus. Ejaculatory duct inserted dorsally represented in New Zealand, is placed in the subfamily inside proximal arms of heavily sclerotized basal sclerite. Molytinae. Female. Sternite 8 with 2 sclerotised arms, sometimes Larvae of Hadramphus and Lyperobius can be separa- fused apically, and bearing short latero-apical setae (Fig. ted from the superficially similar large larvae of Anagotus 70-76). Hemisternites long, divided into distinct proximal species (Aterpinae) by (a) having dorsoepicranial setae 3 and distal portions, the distal portion with numerous short situated on the epicranium and not within the frontal setae; short to long membranous hemisternal pouches suture or on the frons, and (b) abdominal segment IX present. Vagina not extending past hemisternal bases. having two dorsal setae, one slightly longer than the other. Bursa copulatrix large, long; spermathecal duct inserted They are unlikely to be confused, though, because of host- on base of bursa near junction with median oviduct, plant differences — no Anagotus larvae are found on Api- dilated at insertion point. Spermatheca falciform, much aceae, Araliaceae, or Pittosporaceae. shorter than large spermathecal gland (Fig. 77-83). Hadramphus and Lyperobius larvae differ from those of the xylophagous Holarctic Hylobius Germar and Nearctic rv (see, e.g., May 1993, fig. 830-839). Length 17.5- Pachylobius Le Comte mainly in the shape and alignment 32 mm, width 6.5-10 mm. Body large, robust, dense of the spiracles. They share with Hylobius and its allies creamy white; head dark red-brown to black, partially conspicuously large airtubes that are often pigmented. retracted. Frontal suture narrow, not angulate before apex. Larvae of the European molytine Liparus germanus Lin- Dorsoepicranial setae 3 not in contact with frontal suture. naeus are very similar also to those of the New Zealand Frontal setae 5 well developed. Thoracic segments with a Molytini, having elongate-ovate spiracles in common with V-shaped pocket medially between the 2 ventral folds. Hadramphus larvae (cf. Scherf 1964 and May 1971, 1981, Abdominal segment III without prodorsal setae. Spir- 1987a, 1993). acles conspicuous, elongate-ovate or subcircular; airtubes conspicuous, with fine, pigmented annuli, directed dorsad (thoracic) or caudad (abdominal); Abd VIII spiracle on MO O OGICA C A AC E S dorsum, as large as thoracic spiracle. Morphological terms useful in taxonomy and identification are illustrated for external features (Fig. 11-13), male gen- p Maximum length 17-20 mm. Cuticle glabrous; italia (Fig. 62), and female genitalia (Fig. 78). setae dark, tapering, mounted on small tubercles, strongest A character-based phylogenetic species concept (Baum on head and pronotum, finer on abdomen. Elbows with & Donoghue 1995) is adopted in the context of this study. subequal setae. Pseudocerci on abdominal segment I It defines a species as the smallest aggregation of popu- short, dark, curved, without associated setae. Antennal lations diagnosable by a unique combination of character club smooth. Primary protheca bullate (Hadramphus) or states in comparable individuals. striate (Lyperobius), with spinules on dorsal margin and ridges; secondary pterotheca shorter than the primary. d The globose head is produced into a rostrum that [Descriptions of larvae and pupae from May 1971, 1981, bears the mouthparts, comprising a triangular pair of bi- 1987a, 1993.] dentate mandibles with tufted or scattered setae on their outer face, paired maxillae each with a three-segmented t pl nt Apiaceae (Aciphylla and Anisotome), Arali- maxillary palp, and a rectangular mentum bearing a pair of aceae (Stilbocarpa), and Pittosporaceae. three-segmented labial palps. There is variation between the species in the relative length, thickness, and shape of r Pascoe (1876, 1 77 placed Lyperobius huttoni the rostrum. It can vary from relatively short and thick with and L. tuberculatus in Lacordaire's (1863) group Moly- an expanded apex (Fig. 14, 15) to relatively long and thin tides, based on Schoenherr's (1823) family-group name with the apical portion not much more expanded than the for this taxon. The family-group name Liparides (Latreille base (Fig. 16, 17). A dorsal median carina occurs in some 1828) - used by Marshall (1932), Solari (1941), Mori- taxa. The geniculate antennae, inserted laterally on the api- moto (1962), and others to refer to a tribe (Liparini) of the cal third of the cylindrical rostrum, are eleven segmented subfamily Hylobiinae- is equivalent. An ecophyletic cline with the terminal three segments forming a compact club. from fully flying Hylobinii to flightless Liparini was noted The first segment, the scape, contributes almost half the by Morimoto (1982), who associated these two tribes antennal length and is inserted at the anterior end of the under the family-group name Hylobiini Kirby, 1837; the scrobe, a deep lateral sulcus extending obliquely from the tribal name Molytini has priority, however. Following anterolateral portion of the rostrum to the ventral portion
— 9 — of the eye. Segments 2-8 constitute the funicle, and range Sexual dimorphism. Apart from minor differences in in shape from elongate to rounded to transverse. The ratio shape of the rostrum between the sexes, dimorphism is of the length of the first to the second segment, and the most readily observed in the fifth ventrite. The male shape of the 7th funicle segment can be diagnostic of some ventrites are smaller and much more transverse (Fig. 38, taxa. Eyes are lateral, convex, and transverse. 40, 42) than the female ventrites, which are larger, longer, and more rounded on their margins (Fig. 39, 41, 43). Pronotum. Diagnostic features of the pronotum include differences in the degree of subapical and basal con- Genitalia. The male median lobe is stout and robust, as is striction, macro- and microsculpture, and extent of scaling characteristic of molytine weevils. Size and shape of the patterns and punctuation. In Lyperobius the pronotum median lobe and the development of the aedeagal apo- varies from nearly transverse with broadly rounded sides demes relative to the median lobe can be characteristic of to elongate with narrowly rounded sides; such distinctions a particular species. The shape of the endophallic basal are characteristic of particular species and species-groups. sclerite, and the presence or absence of sclerites in the Pronotal sculpture and scale patterning vary from three endophallus apex, can be useful diagnostic characters narrow, longitudinal scale-filled sulci (two marginal, one (Fig. 48-69). In the female genitalia size and shape of the median) (Fig. 21, 24) through the pale scales being con- 8th sternite are useful features for distinguishing species fined to broader, shallower impressions (Fig. 19, 20) to (Fig. 70-76). Degree of development of the hemisternal being scattered in no particular pattern (Fig. 22, 23). In the pouches, size and shape of the hemisternites and styli, and four Hadramphus species the pronotal sides and disc are the nature of their setation can vary greatly between ornamented with tubercles which in size, shape, and posi- species (Fig. 77-83). tion are specific to species or species-groups. The ventro- anterior lateral margins of the prothorax bear conspicuous ocular lobes which partly conceal the eyes in Hadram- phus; these are reduced in Lyperobius. I E IS O Y A IO OGY Larvae of Lyperobius and of two Hadramphus species Elytra. Elytral shape, scale patterns, and modifications of (H. spinipennis, H. tuberculatus) feed on herbaceous per- the elytral intervals are all important taxonomic char- ennial Apiaceae in the predominantly New Zealand genera acters. The degree of prominence of elytral humeri, and the Aciphylla andAnisotome. Fouveaux Strait to Snares popu- development and shape of discal and declivity tubercles, lations of the southern species H. stilbocarpae are found differ greatly between the Hadramphus species (Fig. 1-4). on the fleshy herb Stilbocarpa (Araliaceae), but south- The extent and development of raised costae on the even west Fiordland populations use Anisotome lyallii as a host. and uneven intervals offer a number of useful distin- Larvae are found both in the thick main roots and guishing features in Lyperobius (Fig. 5-10, 27, 28). The rhizomes of the host plants (Bull 1967, May 1971, 1981) shape of the pale scales can vary considerably on a single and in soil surrounding the roots (Bull 1967). Larvae of L. individual, but scale shape is usually relatively constant on hudsoni and L. montanus have also been found grazing on the elytral disc, and can serve as a useful diagnostic leaf bases and crowns. L. huttoni larvae construct a bare character. pupal chamber in the soil adjacent to the roots of the host plant. The pupal stage lasts about 2 weeks. After eclosion Legs. The legs can vary from elongate and slender with the teneral adults remain in the chamber for almost 8 the femora barely claviform and the tibiae straight (e.g., H. months before emerging. The lifespan of adult L. huttoni spinipennis) to short and thick with the femora gradually in the wild has been estimated at 2.5 years (Bull 1967). claviform (e.g., L. cupiendus) and the tibiae short and Larvae of H. pittospori are subcortical borers in the live sinuate (e.g., L. fallax). The extent and position of scaling branches, trunks and root crowns of the woody host plant on the upper inner face of the coxae is an important karo (Pittosporum crassifolium). Their egg-shaped pupal character diagnostic of the L. coxalis species-group (Fig. chambers are found in the centre of the workings. The 32-34). Size and shape of the second and third tarsal pupal stage lasts about 3 weeks. Adults of this species have segments, and the degree of development of the tarsal sole remained alive in captivity for 3 years (Bennett 1987). hairs, offer useful diagnostic characters for distinguishing Hadramphus adults are nocturnal, whereas those of both sympatric congeners and vicarious sister taxa (Fig. Lyperobius are diurnal, feeding actively on leaf tissue, 36, 37). flower stems, and developing seeds (Barratt 1985, Bennett 1987, Bull 1967, Meads 1990). Adult L. huttoni employ
-1 0— two types of feeding — leaf nicking on the outer leaflets of distribution) and the New Zealand Molytini, both groups the host, and a deep ovoid notching of the leaf petioles being predominantly insular or montane to subalpine, with (Bull 1967). Kuschel (1971: 242) described feeding be- many species feeding on plants belonging to or associated haviour of H. stilbocarpae as "adults notch leaf margins at with the family Apiaceae. night." D.S. Homing (in litt.), who observed this species in the field, notes: "We saw many weevils feeding on Stil- Monophyly of in-group clade Hadramphus +Lyperobius. bocarpa and not one of them was notching the leaf margin. Hadramphus and Lyperobius are inferred to be a mono- Rather, they fed on the leaf by chewing a hole through it phyletic clade on the basis of the following combination of and then enlarging the hole. Often this is close to the leaf character states. margin and the narrow strip between the edge of the (1) Spermatheca falciform, with no marked division into feeding hole and the leaf margin may break, creating an nodulus and ramus. Outgroup taxa have a much more illusion that the leaf margin has been notched." Meads' elongate spermatheca with distinctive nodulus and ramus. (1990: 49) photograph of this species feeding confirms (2) Hemisternites divided into distinct distal and proximal Horning's account. Adults of this species can congregate plates (Lyperobius, Fig. 77-78; Hadramphus, Kuschel in large numbers on a single host plant and feed until it dies 1971, 1987). This condition is not found in any out-group (Close 1997). taxa examined, although a somewhat similar state has been noted in the very distantly related Cholinae of the genus Homalinotus (Vaurie 1973, fig. 92, 94) and Brachyderinae Y OGE E IC A A YSIS of genus Cneorhinus (Alonso-Zarazaga 1988, fig. 1). Methods. An attempt is made to reconstruct the phylo- (3) Apical margin of male 8th sternite deeply emarginate genetic history of Hadramphus and Lyperobius using the medially (Hadramphus, Fig. 44, 46; Lyperobius, Kuschel methods of Hennig (1966), which group taxa into mono- 1987). The predominant state in the outgroup taxa and phyletic lineages (clades) on the basis of their possession other curculionid subfamilies is for the apical margin to be of shared derived character states (synapomorphies). either rounded or acute. Character state polarities were determined by outgroup (4) Endophallus of male genitalia non-eversible, attached analysis as discussed by Watrous & Wheeler (1981), Mad- to apical margin of median lobe and contained entirely dison et al. (1984), and Bryant (1991). When two or more inside median lobe. According to Kuschel (1987) this con- states of a character occur within an ingroup, the state dition has not been noted before in curculionids. occurring in related outgroups is considered to be ances- tral (e.g., characters 12, 13, and 14 in the following list). Character list and discussion of character state polarity Equivocal polarity decisions can occur when out-group [0 = ancestral; 1, 2, & 3 = derived] relationships are poorly resolved and uncertain, such as in this study, and two or more states of a character occur both 1. Rostrum median carina absent (0) within and without the ingroup (e.g., characters 5, 7, and 8 Rostrum median carina present (1) in the following list). In these instances the predominant- Hadramphus spinipennis and H. stilbocarpae have a low, states method has been applied, i.e., the state occurring median rostral carina. No such carina is found in most most commonly amongst the outgroup taxa is taken to be outgroup taxa examined. A distinct median carina occurs ancestral for the ingroup (Kluge & Farris 1969). in the distantly related Rhineilipus cuvieri Boheman and A representative selection of phenetically related taxa Plinthus megeleri, and traces of this structure occur in some species of Heilipus belonging to the tribe Molytini, and related genera from (e.g., H. lactarius Germar). the Pacific Rim and Northern Hemisphere, were selected 2. Postocular lobes obsolete (0) as outgroups; they are listed in Appendix 1. All these taxa Postocular lobes well developed (1) were examined for external characters. Genital characters All Hadramphus species have well developed postocular were studied in detail for four taxa — Liparus, Plethes, lobes partly covering the eyes. Postocular lobes are absent Aclees, and Hesychobius. These taxa were selected be- to obsolete in phenetic and phylogenetic relatives (e.g., cause they have either a close phenetic resemblance to Lyperobius, Plethes, Liparus, Anisorrhynchus, Aclees, Lyperobius (Liparus and Plethes) or they are some of the Hesychobius) but variably present in other molytine gen- closest relatives in a geographical sense (Aclees and Hesy- era (e.g., Dyscerus, Apterylobius, Heilipodus). chobius are both found in the southwest Pacific region) to 3. Pronotum lateral margins without tubercles but with a New Zealand taxa. There is also a host and habitat resem- constriction at apical third (0) blance between Liparus (see Magnano 1954 for details of Pronotum lateral margins tuberculate at apical third (1) Pronotum lateral margins tuberculate at middle, and lineatus (Eremninae), the New Zealand Anagotus hamil- widest at middle to apical third (2) toni (Aterpinae), and the South American Astyage linei- In Lyperobius and many other Molytini the pronotum is gera (Cholinae). Amongst the immediate outgroup taxa slightly constricted laterally at the apical third to quarter. only Plethes albolineata has scale stripes resembling Hadramphus pittospori has a large, acutely angled tub- those of Lyperobius. In L. huttoni and L. montanus scale ercle in this position and a small, obtusely angled tubercle stripes are virtually absent on the elytral disc, occurring at the middle (Fig. 1). The other three Hadramphus spe- only on the elytral declivity in a very reduced state in fresh cies possess a large lateral tubercle at the middle of the specimens. pronotal margins (Fig. 2-4). The Chilean Calvertius tub- 8. Elytral disc intervals (3,5,7) flat to convex (0) erosus (Fairmaire & Germain) has a large, protuberant Elytral disc intervals (3,5,7) tuberculate (1) swelling midway on the pronotal lateral margins, but this Elytral disc intervals (3,5,7) costate, with no develop- does not appear to be homologous with the tubercles found ment of secondary costae in even intervals (2) in Hadramphus. Elytral disc intervals (3,5,7) costate, with a tendency 4. Pronotal disc punctate (0) for secondary costae to occur in even intervals (3) Pronotal disc with knobs/tubercles (1) Tubercules on the elytral disc do not occur often in the The pronotal disc varies from sparsely to densely to Molytini and related tribes (exceptions: South American wrinkled punctate in most molytine taxa. Some South genera Byzes, Epistrophus, Placeilipus, Rhineilipus, and American genera (e.g., Byzes, Parabyzes) and Asian Syphorbus, and Plagiophelus from Asia). Such tubercles, species (e.g., Dyscerus perforatus) have their pronotal formed by irregular elevations of the elytral intervals, are disc ornamented with large glossy granules that in some found in all Hadramphus species (Fig. 1-4). Development taxa (e.g., Helipodus onychinus) are coalesced into raised of elytral intervals as raised costae is not a common areas. Only the four Hadramphus species possess four character, occurring only sporadically in a variety of distinct discal knobs/tubercles (Fig. 1-4). curculionid subfamilies, e.g., Astyage lineigera (Cholinae), Lixus lineatus (Cleoninae), several Anagotus species 5. Metasternal margin in front of hind coxae with a (Aterpinae), and several Sargon species (Brachyderinae). metasternal fold, and sometimes a slight overhang at Amongst the immediate outgroup taxa costate elytral most (0) intervals are found in Plethes albolineata and P. ungui- Metasternal margin in front of hind coxae with a culatus. Only the uneven elytral intervals are raised as distinct projecting tooth (1) costae in Lyperobius glacialis, L. carinatus, L. fallax, L. All four Hadramphus species possess a projecting tooth eylesi, and the L. hudsoni species-group (e.g., Fig. 6, 8, 10, on the hind margin of the metasternum directly in front of 27). There is a tendency for the even elytral intervals to be the hind coxae (Fig. 29). Lyperobius and most other moly- raised as secondary costae - though not all individuals in tine taxa have at most a well developed metasternal fold a population manifest this character state — in the L. with a slight overhang (Fig. 30, 31). Elsewhere in the spedenii species-group and in L. australis, L. clarkei, and Molytini and related groups a projecting metasternal tooth L. nesidiotes (Fig. 5, 9). has been noted only in some species of Orthorhinus (e.g., O. aethiops) and Byzes (e.g., B. diversus). 9. Mid elytral declivity with interval 3 flat to costate (0) Mid elytral declivity with a tubercle on interval 3 (1) 6. Mesepisternum with no distinct impression, at most a Both Hadramphus spinipennis and H. stilbocarpae have a few confluent punctures (0) distinctive conical tubercle about midway on interval 3 of Mesepisternum with a distinct elongate impression (1) the elytral declivity. This tubercle has not been observed in The mesepisternum in Hadramphus bears a distinct elon- other Molytini. gate impression which is often filled with pale scales. This impression has not been observed in other Molytini, which 10. Elytral declivity lateral margin without a tubercle on at most have a few confluent punctures forming a very interval 9 just above articulation of abdominal vent- small impression. rites 4 and 5 (0) Elytral declivity lateral margin with a tubercle on 7. Elytral scales scattered (0) interval 9 just above articulation of abdominal vent- Elytral scales confined to even intervals and forming rites 4 and 5 (1) pale scale stripes (1) Hadramphus pittospori, H. spinipennis, and H. stilbocarpae Pale scale stripes on at least some of the elytral intervals have a distinctive tubercle on elytral interval 9 just above are found in a variety of weevil taxa belonging to several the articulation of abdominal ventrites 4 and 5. This tub- curculionid subfamilies, e.g., the South African Eremnus
— 12 — ercle does not occur in H. tuberculatus, Lyperobius, and In most genera of Molytini the apical portion of the other Molytini. endophallus beyond the ostium is completely membranous, 11. Tarsal segments 1 and 2 with apical sole hairs same without any distinct sclerotisations. In Hadramphus (Fig. length as those on segment 3, usually not distally pro- 48-51) and all Lyperobius species except for those of the jected (0) L. coxalis and L. spedenii species-groups (Fig. 55, 56, 66- Tarsal segments 1 and 2 with apical sole hairs longer 68) very distinct sclerotisations are located in the apical than those on segment 3, projected distally (1) portion of the endophallus. All members of the L. hudsoni species-group have the 17. Female styli apical (0) apical tarsal sole hairs on segments 1 and 2 longer than Female styli latero-apical (1) those on segment 3, and they are projected distally (Fig. In L. carinatus (Fig. 78) and L. clarkei the styli are in- 37). This character state is unique to these taxa as com- serted latero-apically on the gonocoxites. In other Lyper- pared to other Lyperobius species (Fig. 36), Hadramphus, obius species (Fig. 77, 79-83), Hadramphus (Kuschel Plethes, and Liparus. 1971, 1987), Liparus, Hylobius, and Pissodes the styli are 12. Femoral tooth or denticle present (0) inserted apically. Femoral tooth or denticle absent, represented at most 18. Female gonocoxites without pouches (0) by a low carina (1) Female gonocoxites with short pouches (1) A tooth or denticle is present on the femora of most species Female gonocoxites with long pouches (2) of Molytini. In the New Zealand taxa it is found only in H. Distinct membranous pouches are located on the proximal pittospori and H. spinipennis. portion of the female gonocoxites of all Hadramphus and Lyperobius species. Two character states, short (e.g., Fig. 13. Coxae with upper inner face bearing scattered hairs 77, 79-83) and long (Fig. 78; Kuschel 1971, 1987), are and scales (0) recognised. These pouches have not been observed in Coxae with upper inner face bearing a dense scale other Molytini. patch (1) The three members of the L. coxalis species-group share 19. Female bursa copulatrix without lobes near junction the synapomorphy of a dense, slightly raised scale patch of oviduct (0) on the upper inner face of all coxae (Fig. 33). This char- Female bursa copulatrix with lobes near junction of acter state has not been observed in any other Molytini oviduct (1) (e.g., Fig. 32, 34). Members of the L. spedenii species-group are character- 14. Male 5th ventrite not medio-apically emarginate (0) ised by a lateral pair of lobes on the bursa copulatrix near Male 5th ventrite medio-apically emarginate (1) the junction with the median oviduct (Fig. 79, 81, 82). This There is a distinct shallow apical median emargination on character state has not been observed in other Lyperobius the male 5th ventrite of H. stilbocarpae and H. spinipennis species, Hadramphus, Liparus, Hylobius, and Pissodes. (Fig. 38). This character state has not been observed in the 20. Female bursa copulatrix completely membranous (0) other Hadramphus species, Lyperobius (Fig. 40, 42), Female bursa copulatrix with distinctive sclerotised Plethes, Liparus, Aclees, and Hesychobius. area near junction with oviduct (1) 15. Female 5th ventrite without an upturned, rimmed All members of the L. hudsoni species-group and L. eylesi outer medio-apical margin (0) possess a distinctive female genital character - a distinct Female 5th ventrite with an upturned, rimmed outer internal sclerite at the junction of the bursa copulatrix and medio-apical margin (1) the median oviduct (Fig. 77, 80, 83). This sclerite is not The medio-apical margin of the female 5th ventrite has an found in other Lyperobius species, Hadramphus, Liparus, upturned rim (Fig. 43) in members of the L. coxalis and L. and Hylobius. An internal female genital sclerite in the spedenii species-groups and in L. eylesi. This character same position has been observed in Pissodes piceae, but it state has not been observed in other Lyperobius species is of a different form to that found in the L. hudsoni (e.g., Fig. 41), Hadramphus (e.g., Fig. 39), and other species-group and L. eylesi. Molytini. Phylogenetic results and relationships. A data matrix 16. Male median lobe endophallus floor at apex beyond was constructed for 15 Hadramphus and Lyperobius taxa ostium without distinct sclerotisations (0) and an hypothetical outgroup taxon based on 73 species of Male median lobe endophallus floor at apex beyond Molytini and related genera, embodying 20 characters ostium with distinct sclerotisations (1)
—13— O t r p p tt p r M j r t r l p tt p r p n p nn 00 8 p n p nn 8 2 00 t lb rp 00 t lb rp t b r l t t b r l t h tt n h tt n l r l r 00 00 r n t r n t l l 00 l l f ll x 4 f ll x
00 h d n pp h d n pp 48 p d n pp 0 0 00 l x l 0 0 p d n pp n d t 0 0 66 C tr l x l 4 l 0 n d t 0 O t r p tr l xt-f 1 M j r t r l n n tr f r a am us nd y e o ius. xt-f lt f b t tr p t t t l t t nd d nd (b l h b r l n br n h nd t th p r nt f th 1 t p r n - br n h pp d nt j r t r l n n tr h p r nt f tr n h h h br n h rr d b t tr p pl n h h h n ph l t r p r pr nt d h n b v h br n h •
Ev l t n 2000 tr pl d r nd l fr th t f ll p bl tr
r r nr t d St rt n d = d = 88
e - ig 3 r n d tr b t n f l n th r n d tr b t n f l n th f 2000 r nd tr : f nr t d r nd tr
0 (6 ( 2 (6 ( 4 ( 0 Ν ( 6 ( • (2 8 ( 2 - _ (4 60 (46 6 (64 62 ( 7 6 (8 64 ( 4 6 ( 79 66 6 68 6 0 2 i W (i
n=66 22000 d=4 68 48 =- 68 8 2= 06 62 using the program AutoDecay 3.03 (Eriksson, n.d.). a e 1 Ch r t r t t d tr b t n tr x f r Analysis of this matrix produced 16 most parsimonious n r l ph l n t n l o a am us trees, with a length of 31 steps. A majority rule consensus nd y e o ius. tree is presented as a summary of phylogenetic relation- a a C a ac e s a es ships (Text-fig. 1). th genera form monophyletic clades, and species relationships are unequivocal in Hadramphus. O t r p A decay index of 5 for this clade (Text-fig. 2) indicates that . i os o i 11111 1 1 1 the internal node uniting all Hadramphus species into one . s i i e is 11 111 111 1 1 clade has greatest support, since a tree of length 37 is the . s i oca ae 11 111 111 1 1 1 shortest without this branch. More problematic are the . u e cu a us 1 111 1 1 1 1 interrelationships of Lyperobius species, but there is only . aus a is 1 3 11 1 ? 1 one unresolved trichotomy in the consensus tree. . ca i a us 1 1 11 . c a kei 13 1 11 S a is ica t t f ph l n t tr t r A bootstrap . co a is 1 11 1 1 test can be used to put confidence intervals on any estimate . ey esi 1 1 11 1 1 of phylogeny by resampling the original data set to infer . a a 1 1 1 1 the variability of the estimate. Characters in a matrix of . g acia is 1 1 1 taxa x characters can be sampled with replacement (i.e., . u so i pp 1 11 1 1 1 bootstrapped) to create many new matrices of the same . u o i 1 1 1 size as the original. Each of these can then be analysed to . esi io es 13 11 1 1 find the best fitting tree. Results can then be combined in . s e e ii pp 13 1 1 11 a majority rule consensus tree. Next to each branch of that tree is shown the number of times that the monophyletic group defined by that branch occurred (Felsenstein 1985). (Table 1). Apart from the outgroup taxon, two other con- t tr p proportions>70% usually correspond to a prob- solidated taxa were used for character state scoring. These ability of >95% that the corresponding clade is real (Hillis were L. hudsoni spp. (comprising L. barbarae, L. hudsoni, & Bull 1993). L. montanus, and L. townsendi) and L. spedenii spp. (L. Judging by published studies, it is frequent for putative cupiendus, L. patricki, and L. spedenii). This matrix was monophyletic groups to be found to be supported with less analysed in the numerical phylogenetics package PAUP than 95% confidence. This is because utility of this test is ver. 3.1.1 (Swofford 1993) using the branch and bound limited in data sets lacking large numbers of characters algorithm, which guarantees finding all the most parsi- relative to the number of taxa (Sanderson 1989). It has monious trees. Decay indices, which indicate the length of been estimated that even in a data set with no character the shortest tree lacking a particular node, were calculated conflict (i.e., homoplasy), every clade must be supported —15— by at least three derived characters for statistical signi- • p tt p r ficance at the 95% level to result. Thus, even under ideal circumstances, where character conflict is low or non- σ Η s i i e is existent, phylogenetic analyses with only a moderate ee e g + C 3 Η t lb rp number of characters per taxon will exhibit low confid- ι 7 ence levels in at least some parts of the resulting tree C 5 Mi 5 σ t b r l t hypothesis (Penny & O'Kelly 1991). M x Application of this test to the reported data (Text-fig. 2) σ h tt n reveals that Hadramphus species form an extremely well supported clade (98%). The most strongly supported clade l r within this group is the sister-group relationship H. spini- pennis-H. stilbocarpae (85%). Support for a Lyperobius σ r n t clade is more equivocal (57%), but within this genus a l l clade comprising the . spedenii and L. coxalis species- groups is better supported (66%). σ f ll x An alternative test, tree-length distribution skewness, can discriminate between character data sets containing σ h d n pp phylogenetic signal and those with significant amounts of random noise. Data matrices consistent with one phylo- σ s e e ii s genetic hypothesis produce tree-length distributions that h t pl nt σ x l are highly skewed to the left, while those compatible with t p many phylogenetic hypotheses produce more symmetrical n rd r d σ n d t tree-length distributions that cannot be distinguished from ΕΞ n n p f those produced by random character data. Skewness of a Ap σ tr l tree-length distribution can be measured by the g 1 stat- Ar l tt p r istic, which is one measure of the skewness of a frequency σ l distribution. Tree-length distribution frequencies that are r ι n rt n significantly skewed at the >99% level (g 1 < —0.47) pro- I v l σ O t r p vide a critical value for distinguishing data sets containing phylogenetic structure from those with random noise e - ig is i u io o os - a ami ies ma e o o (Huelsenbeck 1991). e ma o i y u e co se sus ee o a am us a Applying this test to the reported data set resulted in a y e o ius. tree-length frequency distribution with a mean tree length of 66.7 steps, and highly skewed to the left (Text-fig. 3). tional ranges (expressed as broad habitat groupings, e.g., The g 1 statistic value for this simulation is -1.168, well lowland, subalpine) are mapped onto the independently below the significant critical values reported for this derived phylogenetic tree using MacClade (Maddison & method (Hillis & Huelsenbeck 1992). This value indicates Maddison 1992), and ancestral patterns are predicted by strong support for there being phylogenetic signal in the specifying states for the interior nodes of the tree. This is Hadramphus-Lyperobius data set. equivalent to character state optimisation.
Plant-host analysis. All Lyperobius species utilise only members of the Apiaceae as hosts. The most parsimonious interpretation for the postulated history of host shifts in IS O ICA ECO OGY A IOGEOG A Y the Hadramphus clade involve a shift onto Pittosporum Me o s Biogeographical, ecological, and evolutionary (Pittosporaceae) for H. pittospori, and onto Stilbocarpa history in a phytophagous weevil clade can be estimated (Araliaceae) for H. stilbocarpae from ancestral apiaceous using a phylogenetic tree derived from morphological or host plants (Text-fig. 4). other evidence. This reconstruction of phylogeny gives a Speciation and the evolution of host-plant shifts in critical basis for postulating scenarios of host-plant usage, extant species of Hadramphus and Lyperobius happened selection, and shifts (Anderson 1993), as well as other well after the divergence of the higher taxa of the plants on aspects of biogeographic and ecological history (Brown which they feed. These host-plant shifts have occurred 1995, Morrone & Crisci 1995). Host plants and eleva- among plants that appear very similar in anatomical,
-1 — chemical, and embryological features (Harbourne & Table 2 Numbers of ancestral nodes in the phylogenetic Turner 1984, Hegnauer 1969, Metcalfe & Chalk 1950). In tree separating vicariant and sympatric clades. an analysis of DNA sequences Apiaceae was the closest Clade Ancestral nodes (n) relative of Araliaceae, and this clade was the sister taxon to Pittosporaceae (Chase et al. 1993). Host-plant relations of (1) Vicariant New Zealand Molytini offer some support for the H. spinipennis-H.stilbocarpae 1 hypothesis that chemical similarity and phylogenetic H. pittospori-H. spinipennis/H. stilbocarpae 1 affinity of insects' host plants facilitate host shifts. u e cu a us- i os o i/Η s i i e is/ Plant hosts for most non-New Zealand Molytini are a stilbocarpae 1 wide variety of gymnosperm and herbaceous or woody L. carinatus-L. clarkei 1 angiosperm taxa (Anderson 1993). They include Coni- L.glacialis-all th r Lyperobius 2 ferae for the northern temperate genera Hylobius, Hylo- L. fallax-L. hudsoni pp / eylesi/L. spedenii pp / bitelus, Pissodes, and Pachylobius, Araucariaceae for the coxalis pp 1 Chilean Calvertius, Ficus (Moraceae) for the Asia-Pacific L. coxalis-L. australis/L. nesidiotes 1 Aclees, and genera of Aceraceae, Oleaceae, Rosaceae, Illi- L. australis-L. nesidiotes 1 ciaceae, and Lauraceae for the Asian Dyscerus (Morimoto 1982). Some plant hosts for genera that are close phenet- ( S p tr H. stilbocarpae-L. nesidiotes 12 ically to Hadramphus and Lyperobius include Asteraceae L. carinatus-L. huttoni 3 and Apiaceae for the Palearctic Liparus (Smreczynski L. carinatus-L. spedenii 6 1968), and the South American Helipodus (rearing label L. clarkei-L. fallax 3 data in NZAC), and Asteraceae for the northern Andean L. carinatus-L. fallax 3 Plethes (rearing label data in NZAC). This crude 'out- L. huttoni-L. fallax 3 group' comparison provides additional support for the L. hudsoni pp - spedenii pp 3 hypothesis, derived from phylogenetic analysis, that the ancestral host plant for the Hadramphus-Lyperobius clade was a member of the Apiaceae. spedenii are also vicariant, the former species occurring on A review of specialised associations between insects North Otago and South Canterbury mountains and ranges, and Apiaceae (Berenbaum 1990) noted that records of the latter on Central Otago and Otago Lakes mountains apiaceous feeders or their relatives on araliaceous hosts and ranges and the Takitimu Mountains. Other vicariant were quite rare. Besides Molytini, another New Zealand patterns of disjunction include the sister species L. clarkei curculionid group (Stephanorhynchus, Eugnominae) has (northwest Nelson and Buller ranges) and L. carinatus species-groups host specific to Apiaceae, Araliaceae, Pit- (Seaward Kaikoura Range to mid Canterbury), and the tosporaceae, or Asteraceae (Hudson 1934, Styles 1973, sister-group relationship of L. fallax (northern South May 1987b). Species-groups of Lepidoptera in Xyridacma Island ranges, with southern limits north of the Rakaia (Geometridae), Acrocerops (Gracillaridae), and River) to a clade comprising L. coxalis, L. hudsoni, and L. Pterophorus (Pterophoridae) are found on Araliaceae and spedenii species-groups plus L. eylesi (southern South Pittosporaceae, with distinct species associated with each Island and The Snares, with L. spedenii extending north- family (Dugdale 1975). wards to just south of the Rakaia River) (Craw 1990). Vicariant distributions are most common between Vicariant distribution and sympatric occurrences. The closely related clades and species, with an average of 1.12 major clades Hadramphus and Lyperobius are almost tot- ancestral nodes separating vicariant taxa in the phylo- ally vicariant, and their distributions overlap only at the genetic tree. Sympatric occurrences are usually between southernmost limit of distribution, on tiny Broughton much more distantly related taxa, with an average of 4.7 Island in the subantarctic Snares group. These two taxa do ancestral nodes separating those taxa that occur together occur in geographical proximity in Canterbury, but exhibit (Table 2). Sympatry, due to dispersal, is thus secondary to ecological vicariance along an altitudinal gradient, with H. vicariance in these two genera. tuberculatus known from lower altitudes than Lyperobius species. Elevational range, habitat, and biogeographic history. Members of the L. hudsoni species-group (L. barbarae, Tectonic and other geological activity tends to result in L. hudsoni, L. montanus, L. townsendi) and the L. coxalis irregular land surfaces. These irregularities promote bio- species-group (L. australis, L. coxalis, L. nesidiotes) are logical diversification by changing environments and sep- all vicariant. Putative closest relatives L. patricki and L. arating or joining habitats. Tectonic processes leading to
— 17— uplift, subsidence, and horizontal displacement along fault is reported from South Westland (Sutherland 1996). In zones are postulated to be responsible for the many parts of New Zealand the uplands form disjointed fragmentation of ancestral New Zealand molytine and faulted blocks, remnants of ancient coastal plains, populations, leading to vicariant differentiation at generic which have been uplifted to their present position. and specific levels. Evolutionary and ecological differentiation of the alp- Repeated episodes of uplift and subsidence, and marine ine biota is closely linked to these earth history events. transgression and regression, have characterised the geo- Rocky and other coastal, insular, lowland, and alpine logical history of New Zealand since the Mesozoic era. habitats are ecologically similar. Not only are there eco- Mountain ranges have been a prominent feature of local logical correlations between these environments but also landscapes since those times. Published paleogeographic historical ones. Many montane, subalpine, and alpine spe- maps illustrate mountainous landforms in what is now cies have coastal, insular, or lowland disjunct populations southern Nelson/north Westland, east Otago, and Fiord- or vicariant relatives. Coastal, insular, and alpine biota are land during Lower Cretaceous time. It is often stated that ecologically and historically homologous (Heads 1990). the extensive New Zealand microcontinent had been Most Lyperobius species occur in higher montane, sub- eroded to an expansive peneplain surface by the late alpine, and alpine grassland, herbfield, and rocky places in Cretaceous, but the description `peneplain' is problematic. the South Island. Lyperobius huttoni has a wider tolerance Beneath the very late Cretaceous Pakawau coal measures in elevational range than other species, with populations at surface relief was at least 100 mover very short distances. low altitude on the southern Wellington coast, montane in Similar relief, though less steep, is known from beneath the Hunters Hills (South Canterbury), and subalpine on Eocene coal measures in southwest Nelson and around several South Island ranges east of the main divide. The Huntly (Waikato). In north Westland, the Cretaceous southernmost species, L. nesidiotes, is confined to a small Porarari Group is thought to have been laid down during a islet in the subantarctic Snares islands. period of intense local tectonic activity, and the source of In contrast, Hadramphus species have a strictly the breccia was probably at a considerable altitude. There coastal, insular, and lowland elevational range (Text-fig. was a general tendency for continuing uplift until the end 5). This genus appears to have been more widely distrib- of the Rangitata Orogeny in the late Cretaceous (Suggate uted in the past, as Dr T. Worthy has recently discovered a et al. 1978). large Hadramphus specimen in a subfossil deposit on the No epoch of the Cenozoic era lacks deposits indicative west coast of the South Island (Te Ana Titi cave, Fox of tectonic movement somewhere in New Zealand. During River, 12 Mar. 1992 - NZAC). No large molytine species the Paleocene to Late Eocene there was a land of generally is known at present to inhabit this area. This subfossil low relief flanked by large, low-lying lagoon areas in parts specimen resembles a large female Hadramphus in NZAC of the eastern South Island, with localised hill and moun- collected at Puysegur Point, and identified below as H. tain ranges. Late Eocene ranges existed to the north and stilbocarpae. In light of this discovery the Puysegur Point south of the Manapouri region. Minor mid-Cenozoic population of Hadramphus urgently requires specialist uplift affected the western edge of the shelf region in the study, since it and the subfossil material may represent a southern South Island, from the Longwood Range in the fifth, as yet undescribed, Hadramphus species. south through the Takitimu Mountains to the Bob's Cove It has been suggested that some ancestral alpine plant area (Lake Wakatipu region) in the north. Early Oligocene species of ancient mountains could have survived on and mid Miocene ranges occurred in the Takitimu area. In `rocky faces' or in habitats characterised by cool, wet, Otago, the Moonlight Fault is bordered by a discontinuous infertile old soils at low altitudes during the early to mid narrow strip of Oligocene beds, interpreted as a tectonic Cenozoic, and later given rise to the present alpine flora zone active during Oligocene sedimentation and Kaikoura (Cockayne 1928, Wardle 1968, 1978). Particularly rele- deformation (Norris & Turnball 1993). vant is the fact that numerous species of the probable Initiation of the Kaikoura Orogeny, which gave rise to ancestral host-plant family and genera (Apiaceae: Aci- the present-day axial ranges, can be placed with reason- phylla and Anisotome) occupy coastal cliff and lowland able confidence in Early Miocene time for many parts of peaty and swampy habitats today (Dawson 1971). New Zealand. The modern Southern Alps may have Insular and coastal weediness characterised Mesozoic developed northwards from an embryonic mountain range weevil taxa ancestral to modern forms. The New Zealand in north-west Otago originating through significant late Molytini are hypothesised to have descended from a Late Oligocene - early Miocene uplift (Craw 1995). A distinct Cretaceous - early Tertiary brachypterous ancestral com- period of early Miocene uplift occurred in Marlborough plex distributed in coastal insular and lowland sites, and (Baker & Seward 1996), and middle to late Miocene uplift on mountain ranges. That the common ancestor of Had-
—18— Table 3 Taxonomic root weight values for Hadramphus and Lypero- Table 4 Summed percentage taxonomic bius species and species groups, diversity values (TDV) for selected critical areas where two or more Hadramphus and/ Taxon Information Quotient Weight Percentage or Lyperobius species are sympatric. Areas i os o i 3 24.6 3 8.81 ranked for importance by this measure. s i i e is 4 18.5 2.26 6.64 Area TDV Rank s i oca ae 4 18.5 2.26 6.64 u e cu a us 2 37 4.51 13.2 Black Birch Ra. 19.8 1= u o i 2 37 4.51 13.2 Craigieburn Ra. 19.8 1= c a kei 4 18.5 2.26 6.64 Seaward Kaikoura Ra. 19.8 1=. ca i a us 4 18.5 2.26 6.64 NW Nelson/Buller 13.3 2= g acia is 3 24.6 3 8.81 Broken River Basin Ι 3 3 2= a a 4 18.5 2.26 6.64 Murchison Mtns 13 3 u so i spp. 5 14.8 1.8 5.3 The Snares islands 9.5 4 s e e ii spp. 7 10.6 1.3 3.8 Garvie Mtns 9.1 5= co a is 8 9.25 1.13 3.3 Takitimu Mtns 9.1 5= esi io es 9 8.2 1 2.9 North Otago mtns 9.1 5= aus a is 9 8.2 1 2.9 The Remarkables 9.1 5= ey esi 6 Ι 3 1 5 4.4
σ Η p tt p r ramphus and Lyperobius was flightless is a reasonable hypothesis on the basis of parsimony, since all extant σ Η p n p nn species of these genera are brachypterous and flightless. r l n th + Widespread ancestral populations were subjected to CI 3 t lb rp I 7 phases of `modernization' during which tectonic C 5 displacement, subsidence, and uplift disrupted and t b r l t Μi 5 ® fragmented them, with subsequent differentiation at the M x σ h tt n generic and specific levels.
■ l r
Ει r n t CO SE A IO S A US A A UE Large weevil species are generally considered in New l l Zealand conservation biology (e.g., Ramsay et al. 1988, Thomas 1996) to be at threat from introduced rodent ■ f ll x predators. There is some anecdotal and empirical evidence
• h d n pp supporting this supposition. Bull (1967) provided evi- dence that mice prey upon L. huttoni adults, and Kuschel • p d nhl pp (1971) attributed the absence of H. stilbocarpae on Big South Cape Island - where it was once present - to pred- h b t t x l ation by rats. While rodents undoubtedly prey on these t p n rd r d weevils, other factors such as habitat degradation and n d t n n p f destruction due to agricultural practices, reduction in the n l r size and extent of host plant populations through browsing l tr l ί l l nd by stock and wild goats (Meads 1990), and even removal b lp n 1 l of the entire habitat through quarrying (which happened to •" = n rt n a viable population of L. huttoni at Owhiro Bay, south
® v l O t r p Wellington coast) pose just as great a threat as rodents to populations of these endangered flightless species. Rod- Text-fig. 5 Distribution of habitats mapped onto j r t ents may also compete for host plants with these weevils, r l n n tr f r a am us nd y e o ius. as Roberts (1895) has noted that introduced rats feed on
-19- the roots of Aciphylla. protection, consideration should be given to protecting In 1980 H. spinipennis, H. stilbocarpae, H. tubercu- some of the more speciose high-altitude Lyperobius latus, and L. huttoni were added to the protected fauna list clades. Rather than adding a further early diverging in the Seventh Schedule of the Wildlife Amendment Act species to the protected list, members of the montane to 1980. Being taxonomically and ecologically diverse, indi- subalpine L. coxalis, L. hudsoni, or L. spedenii species- vidually habitat specific, phytophagous on specific host groups could be granted consideration as the next genera, and easily identified, these relatively large weevils conservation priority. Placing priority on one or more of would seem to be an ideal biological indicator group for these speciose clades would also maximise the diversity of the purposes of conservation and environmental habitats occupied by the conserved species. At present assessment. most conservation effort and legislation are focused on There is increasing interest in utilising phylogenetic basal lineages inhabiting coastal, lowland, and insular hypotheses as measures of biodiversity in prioritising, habitats. ranking, and targeting species for `conservation value' (e.g., Williams et al. 1991, 1993, Faith 1994, Humphries eta!. 1995). Application of the taxonomic root weighting method (Williams et al. 1991) to the phylogenetic tree provides a means of assigning quantitative values for each ESC I IO S species or species-group in that tree (Table 3). Fortuitously, root weighting analysis indicates that, Ge us dr ph ou given the present situation and current legislation, pro- Hadramphus Broun, 1911: 104-105. -Hudson 1923: 390. tection of these endangered, rare, and vulnerable species -Schenkling & Marshall 1931: 18. -Hudson 1934: covers 39.68% of total taxonomic diversity of New Zea- 214. -Kuschel 1971: 240. —May 1971: 293. land Molytini. According to this method the next taxon Type species Hadramphus spinipennis Broun, 1911, that should be added to the protected fauna list is either H. by original monotypy. pittospori or L. glacialis. Addition of either one or other Karocolens Kuschel, 1987: 15, new synonymy. of these taxa would bring protected taxonomic diversity Type species Karocolens pittospori Kuschel, 1987, by coverage up to 48.49% of total taxonomic diversity for this original designation. group. Nearly 50% of total taxonomic diversity would therefore be protected by listing only 25% of species in Redescription. Length 11.7-23 mm, width 6.5-11.5 mm. this clade. Since three Hadramphus species are already on Prothorax with ocular lobes well developed and cov- the list, it would be more desirable to conserve L. glacialis ering eyes. Pronotum with 4 discal tubercles and 1 or 2 to maximise the number of clades protected. Root weight lateral tubercles. Elytral intervals 3, 5, and 7 with tub- values can also be combined with information from geo- ercles. Mesepisternum with a distinct elongate impression. graphic distribution, to identify and rank `critical areas' Metasternum in front of hind coxae with a well developed for biodiversity preservation, by summing the percentage projecting tooth. taxonomic diversity values for a selection of areas where two or more species occur (Table 4). Remarks. When describing Karocolens, Kuschel (1987: In setting conservation priorities taxonomic root weigh- 15-16) wrote as follows. ting gives much higher weights to earlier diverging, bas- "Closely related to Hadramphus Broun but differing in ally branching taxa (Humphries et al. 1995). While there the following main characters. are sound reasons for assigning high conservation priority Frons with large impression. 2nd segment of funicle to taxa basal within their lineages (Stiassny & de Pinna roughly as long as 1st. Club segment 1 slender, nearly 1994), weighting methods that emphasise their species- stalked at base, widening in straight line. Prothorax as or rich sister clades also deserve consideration. An alterna- nearly as long as wide, diverging in straight line to anterior tive approach is to attempt to conserve taxa that may be third, with large triangular knob at this point and a small able to support further evolution. This is the methodology one behind. Elytra widening in straight line to anterior advocated by Erwin (1991), who suggests that members of third, with small tubercle or swelling on the basal angles, rapidly evolving clades or `evolutionary fronts' have more with 1 or 2 posthumeral tubercles on interstria 9. Femora potential to lead to future diversity than species-poor armed with tooth; tibiae sinuous on lower edge, with dis- lineages. As basal lineages of New Zealand Molytini are tinct premucro in both sexes. species poor and have already received a much higher "MALE: Tegmen with a pair of teeth on anterior margin weighting in conservation prioritisation and legal of ring and broad parameres. Aedeagus with thick, per-
—20-- — manently lifted ostial valves; floor of internal sac without long, 2.26-2.49x its apical width. Head with a distinct sclerite beyond ostium at aedeagal apex. impression between eyes. Funicle with first 2 segments "FEMALE: Tergal pouches at basal angles of 8th tergite subequal in length. Antennal club with lst segment elon- present, large, bearing filamentous prokaryotes (bacteria). gate, almost stalked at base. Pronotum (including lateral Hemisternal pouches long." tubercles) as long as wide or nearly so, widest at apical Some of the characters listed as distinguishing Karo- third, sparsely granulate, with 4 discal tubercles and a colens, though, are shared with Hadramphus species. For median carina between anterior tubercles. Pronotum lat- instance, H. spinipennis has a femoral tooth that is visible erally with a large triangular knob at apical third and a in specimens without the assistance of any magnification, small median knob. Elytra elongate, widening in a straight and all Hadramphus species have a premucro in both line to apical quarter, with a humeral tubercle; interval 5 sexes. Other characters shared with Hadramphus are pres- with 3 or 4 tubercles on apical three-fifths, interval 7 with ent even though they are stated to be absent in Karocolens. 4 tubercles on apical three-fifths, interval 9 with 1 or 2 In the male genitalia of K. pittospori the endophallus has a posthumeral tubercles and an apical tubercle just above pair of sclerites embedded in the membranous floor, be- articulation of ventrites 4 and 5. Femora armed with a yond the ostium, at the aedeagal apex which appear to be small tooth. homologous with the single inverted V-shaped sclerite Male. Tegmen with a pair of teeth on anterior margin of found in other Hadramphus species. ring; parameres broad (Fig. 45). Aedeagus with a short, Although a number of unique characters (within the stout median lobe and thick, permanently raised ostial clade comprising the New Zealand Molytini) remain that valves; apodemes longer than median lobe. Endophallus phenetically separate H. pittospori from other Hadram- with a pair of sclerites beyond ostium at aedeagal apex; phus species, phylogenetic analysis demonstrates that this basal sclerite heavily sclerotised on anterior half, less species belongs to a clade comprising (H. tuberculatus + sclerotised and with vestiture on posterior half (Fig. 48). (H. pittospori + (H. spinipennis + H. stilbocarpae))) (see Female. Hemisternal pouches long. Spermathecal duct Text-fig. 1). If formal classification is to be as isomorphic not reaching past end of bursa. as possible with reconstructed genealogy (e.g., Darwin 1859, Hennig 1966, Nelson 1973), it is more informative Material examined. Type series only (AMNZ, MONZ, to recognise two genera only in New Zealand Molytini, NZAC). and to synonymise the monotypic genus Karocolens under Hadramphus. Immature stages support this synonymy. Distribution. ND: Poor Knights Islands (Aorangi, Ta- May (1987: 30) notes: "The larva and pupa of K. pittospori whiti Rahi). exhibit only minor difference from the immature stages of H. stilbocarpae and from those of Lyperobius Pascoe." Host plant. Pittosporum crass(folium (Pittosporaceae).
Remarks. This distinctive species, or an as yet unknown sister species, may exist in coastal lowland forest in the dr ph p tt p r (Kusc e ew com i a io North Island and the Marlborough and Nelson areas of the Figures 1, 44, 45, 48; Map 1 South Island, as well as on offshore islands where woody pittospori Kuschel, 1987: 15-18 (Karocolens). -May Pittosporaceae or Araliaceae occur. 1987a: 29-34.
Type data. Holotype: male, ND, Poor Knights Islands, dr ph p n p nn ou Aorangi, 15 December 1981, reared from Pittosporum Figures 2, 49; Map 2 c assί o ium wood collected 1 month earlier, J.C. Watt spinipennis Broun, 1911: 105-106 (Hadramphus). —Hud- (NZAC), 19.7x8.5 mm. son 1923: 390. —Kuschel 1971: 240. Paratypes (6 male, 6 female): listed in Kuschel (1987). Redescription. Length 16.5-19.7 mm, width 7.5-8.5 mm. Type data. Lectotype: male (here designated), Chatham Integument dark brown to blackish brown; body vesti- Islands, "Pitt Is." [T. Hall] (BMNH), 20.6x10 mm. There ture of elliptic to lineal-lanceolate scales, densest on dorsal is one specimen of this species in the Broun Collection surface, the scales yellowish on sides of dorsal surface and (BMNH) and none in the duplicate Broun Collection , reddish-brown to cinnamon on median areas of pronotum (NZAC). The original description indicates that Broun and elytra and on elytral declivity. Rostrum relatively had more than one specimen when he described this spe- cies, hence the lectotype designation. —21— Redescription. Length 20.2-23 mm, width 9.6-11.5 mm. Redescription. Length 15.5-21.7 mm, width 7.8-9.5 mm. Integument dark reddish-brown; body vestiture of pale Integument dark brown; vestiture of greyish-brown to yellowish to dark brown, lineal-lanceolate, appressed dark brown, appressed, lineal-lanceolate scales covering scales and hairs, densest on dorsal surface. Pronotum with body; scales on top of tubercles paler. Pronotum with lateral lines and median apical and basal streaks of pale lateral and median pale lines; lateral lines often extending yellowish scales. Rostrum relatively long, 2.3-2.6x its back onto elytral interval 5. Rostrum relatively long, 2.4- apical width. Funicle with 1st segment 1.5-1.72x as long 2.7× its apical width, usually with a weak median carina or as 2nd segment. Pronotum (including lateral tubercle) smooth line. Funicle with 1st segment 1.4-1.7× as long as wider than long. Elytra oblong-ovate; tubercles on inter- 2nd segment. Pronotum (including lateral tubercles) 1.15- vals 3 and 5 conical, on interval 7 spinose; interval 9 with 1.35× wider than long, with distinct discal knobs; surface a tubercle just above articulation of ventrites 4 and 5; granulose to granular rugose. Elytra elongate ovate, with declivity with a median tubercle on interval 3. surface granulate; interval 7 with conical tubercles, inter- Male genitalia. Median lobe stout; aedeagal apodemes val 9 with a tubercle just above articulation of ventrites 4 approximately as long as median lobe. Endophallus apex and 5; declivity with a median tubercle on interval 3. with an inverted U-shaped sclerite (Fig. 49). Male. Median lobe short, stout; aedeagal apodemes Female genitalia. Sternite 8 elongate; apodeme long. shorter than median lobe. Endophallus apex with an in- Hemisternites with proximal portion noticeably broadened; serted U-shaped sclerite (Fig. 50). hemisternal pouch moderately long. Female. Sternite 8 with apodeme long. Hemisternites with proximal portion noticeably broadened; hemisternal Material examined. Lectotype, plus 21 non-type exam- pouches long. ples (BMNH, NZAC). Material examined. Type series, plus 160+ non-type Distribution. Chatham Is: Mangere I., Pitt I., Rangatira examples (AMNZ, BMNH, CMNZ, MONZ, NZAC, (South East) I. OMNZ).
Host plant. Aciphylla dieffenbachii (Apiaceae). Distribution. FD: Breaksea Is, Breaksea Sound; Reso- lution I. (exposed outer coast and Five Fingers Peninsula); Remarks. The conservation status of H. spinipennis has Puysegur Point. SI: Bird I. (Fouveaux Strait); Big South been evaluated by Emberson & Marris (1994) and Cape I. The Snares (North East I., Broughton I.). Emberson et al. (1996). It would be worth searching for this endangered and legally protected species on other Host plants. Anisotome lyallii (Apiaceae); Stilbocarpa islands in the Chathams archipelago. For instance, it has lyallii, S. o us a (Araliaceae). never been recorded on Rekohu (the main island), but could well occur there on the southern coastal cliffs, as Remarks. Larvae and pupae have been described by May well as at higher elevations on the Southern Tableland, (1971, 1981). Colour photographs of an adult and its hab- where populations of a possible alternative host plant, itat, along with valuable notes on behaviour and ecology, Aciphylla traversii, occur. Population dynamics of this can be found in Meads (1990). In addition, H. stilbocar- species have been studied by Schöps et al. (1998). A pae has been illustrated and discussed by Rance & Patrick colour photograph of this species appears in Dugdale & (1988), Morris & Hayden (1995), Peat & Patrick (1996), Emberson (1996: 97). and Close (1997). Translocation efforts and other aspects of conservation management of this species have been reported by Thomas et al. (1992), Meads (1994), and dr ph t lb rp K h l Thomas (1996). Figures 3, 29, 32, 38, 39, 46, 47, 50; Map 3 stilbocarpae Kuschel, 1971: 240 (Hadramphus). -May dr ph t b r l t ( 1971: 293; - 1981: 269. Figures 4, 51 Map 4 Type data. Holotype: male, SI, Big South Cape Island, tuberculatus Pascoe, 1877: 142 (Lyperobius). -Broun Murderers Cove, 4 January 1955, R.K. Dell & B.A. 1880: 445. -Hudson 1923: 390. -Dalla Torre, Holloway (NZAC), 17x8.7 mm. Schenkling, & Marshall 1932: 66. -Kuschel 1971: Paratypes: listed in Kuschel (1971). 240 (Hadramphus).
- - Type data. Lectotype: male (here designated), MC, the presence of Aci y a species in a variety of habitats, "Christchurch" [C.M. Wakefield] (BMNH), 14.5x7.5 including sandhills, swamps, and plains. These host plants mm. for u e cu a us have been recorded from Hagley Park (near the centre of Christchurch) as late as 1905 (Herriott Redescription. Length 11.7-16.3 mm, width 6.5-8.3 mm. 1919), and the last record of this weevil in Christchurch Integument dark brown; body vestiture of greyish- was in 1910. Distribution data from known specimens brown, appressed, lineal-lanceolate scales, those on head, extends from Waimate and Temuka in eastern South Can- elytral intervals, and top of tubercles paler. Pronotum with terbury to Oxford in southern North Canterbury, with pale lateral and median stripes on basal half. Rostrum western and eastern limits at Mt Oakden and Christchurch relatively long, 2.0-2.16x its apical width. Funicle with respectively. Subfossil records of this species from South 1st segment 1.46-1.8× as long as 2nd segment. Pronotum Canterbury (Worthy 1997) do not represent significant (including lateral tubercles) 1.13-1.39× wider than long, extensions to the known range. with 4 low discal elevations. Elytra oval, widest near middle, with surface granulate; interval 3 with 3 how flat elevations, the basal one almost costate, the others tuber- culate; interval 5 with 2 mid-discal, low, flat, tuberculate elevations; interval 7 with 3 low, flat, tuberculate ele- vations; declivity without tubercles on intervals 3 and 9. Ge us y e o ius ascoe Male. Median lobe stout; aedeagal apodemes a little y e o ius Pascoe, 1876: 54. –Broun 1880: 445.–Hutton shorter than median lobe. Endophallus apex with an 1904: 203. –Hudson 1923: 390. –Dalla Torre, inverted U-shaped sclerite (Fig. 51). Schenkling, & Marshall 1932: 65. –Hudson 1934: Female. Sternite 8 elongate; apodeme long. Hemisternites 133. –Kuschel 1971: 239; — 1987: 15. Type species with proximal portions noticeably broadened; hemisternal y e o ius u o i Pascoe, 1876, by original pouches short. monotypy.
Material examined. Lectotype, plus 16 non-type ex- Redescription. Length 11.5-28 mm, width 6-12.7 mm. amples (BMNH, CMNZ, NZAC). Prothorax with ocular lobes slightly developed. Pro- notum not tuberculate, its sides subparallel to broadly Distribution. NC, MC, SC: Canterbury plains, foothills, rounded, without lateral knobs. Elytra not tuberculate; and fringing ranges – Oxford, Christchurch, Blackford intervals flat, convex, or costate. Mesepisternum with no and Mt Oakden (Rakaia R.), Temuka, Waimate. distinct impression, at most with a few confluent punc- tures. Metasternum in front of hind coxae with at most a Host plants. Probably Aci y a su a e a a and A. prominent overhang. g aucesce s (Apiaceae).
Remarks. This legally protected species is possibly ex- y e o ius aus a is ew s ecies tinct (last specimen collected Waimate, 1922). It is men- Map 5 tioned in important conservation publications as having once occurred on Banks Peninsula (e.g., Wells e a 1983, Type data. Holotype: female, FD, Mt Inaccessible, 3400 Ramsay e a 1988, Tisdall 1994). This distribution infor- ft [1020 m, 5 February 1985, A. Frampton (NZAC), mation is based on Kuschel (1971: 240), who noted " 23x9.7 mm. u e cu a us is known mainly from Banks Peninsula but Paratypes: 1 female, FD, near Lake Fraser, Feb 1983, was found as far south as Waimate." None of the known A.E. Frampton. specimens bears a locality label `Banks Peninsula' nor any label data that could possibly be interpreted as indicating Description. Length 20.5-23 mm, width 9.2-9.7 mm. collection there. The nearest recorded specimens to Banks Integument black to dark reddish-brown; body vestiture Peninsula are from Christchurch, including the type series, sparse, of pale yellowish-white scales, densest on head and of which Pascoe (1877: 142) wrote: "Hab. Christchurch ... pronotum, as narrow stripes between elytral striae and found on a plant called the `Spaniard' [i.e., Aci y a] " costate intervals, and on upper inner face of all coxae. Early accounts of the indigenous vegetation around Rostrum relatively long and thin, 2.0-2.05x its apical Christchurch contemporaneous with the collection of the width. Funicle with 1st segment 1.6-l.66x as long as 2nd type series (e.g., Travers 1869, Armstrong 1870) record segment; segments 3-6 about as long as broad; 7th seg-
— 3— ment transverse. Pronotum transverse, l.2-1.25x wider and 2 with apical sole hairs longer than those on segment than long, with sides broadly rounded; disc with a median, 3, and greatly projecting distally. elongate, low, flat, smooth area and a distinct basal im- Male. Ventrite 5 with a distinct medio-apical impress- pression. Elytra elongate elliptic, with even and uneven ion. Median lobe stout; aedeagal apodemes about as long elytral intervals raised as distinct costae; intervals 3 and 9 as median lobe. Endophallus with basal sclerite bearing costate and conjoint on elytral declivity. Coxae with a lateral apical processes; apex with a pair of sclerites (Fig. dense scale patch on upper inner face. 52). Μ l Un n n Female. Ventrite 5 convex on basal half, with distinct Female. Ventrite 5 very convex in basal half, with a lateral impressions. Sternite 8 elongate; apodeme long medio-apical longitudinal impression and upturned apical (Fig. 70). Hemisternites with abundant long, fine setae on margin. Sternite 8 broad; apodeme long and thin. Hemi- distal portion; hemisternal pouches relatively short to sternites with stout setae on distal portion; hemisternal short. Bursa copulatrix with a distinct, broad, sclerotised pouches short, broad. area near junction with oviduct (Fig. 77).
Material examined. Type specimens only (NZAC). Material examined. Type series, plus 22 non-type ex- amples (BHPC, BIPB, NZAC, OMNZ). Distribution. FD: mountain ranges between Edwardson Distribution. MK, SC, CO, DN: Mt Sutton, Ben Ohau Sound and Dusky Sound (e.g., Kakapo Ra.). Ra. (southern end), St Mary's Ra., Hawkdun Ra., Ida Ra., Kakanui Mtns. Host plant. Probably an A iso ome species (Apiaceae). Host plants. Aci y a o so ii A e uosa A mo a a Remarks. y e o ius aus a is is easily distinguished var. g aci is (Apiaceae). from its closest relatives ( co a is and esi io es by the combination of elytra elongate elliptic, pronotal sides Remarks. y e o ius a a ae is easily distinguised broadly rounded, elytral intervals 3 and 9 costate and from its closest relatives u so i and ow se i by conjoint on declivity, and female 5th ventrite with basal its much less developed, more diffuse and indistinct sulci half very convex. and scale pattern on the pronotal disc (cf. Fig. 18, 21, 24). A colour photograph can be found in Patrick (1991, fig. 9). This distinctive species is named for Dr Barbara Barratt, p r b b rb r ew s ecies whose entomological research in the southern South Is- land mountains has contributed greatly to this revision. Figures 18, 30, 52,70, 77; Map 6 Type data. Holotype: female, MK, Mt Sutton [west of Oh ] ft [ 1 ] 9 v b r 19 Μ Fitzgerald (OMNZ, ex R.C. Craw coll.), 20.8x9.7 mm. p r b r n t ou Paratypes: 2 males, 2 females, MK, Ohau Ski Field, 31 Figures 53, 71, 78; Map 7 Dec 1988, B.H. Patrick. ca i a us Broun, 1881: 702 ( y e o ius -Hutton 1904: 203. —Hudson 1923: 390. —Dalla Torre, Schenkling, & Description. Length 16.7-20.8 mm, width 8.5-9.7 mm. Marshall 1932: 65. -Hudson 1934: 214. Integument black to dark reddish-brown; body vesti- ture of pale whitish-yellow to white, oval to ovate scales, Type data. "No locality, came from J.D. Enys, Esq." densest on lateral margins of pronotum and elytral inter- (CMNZ), 20.4x9.2 mm. vals 2, 4, and 6. Rostrum relatively long and thin, 2.23- This specimen bears two 19th century labels reading 2.45× its apical width. Funicle with lst segment 2x as long "carinatus" [in Broun's hand] and "J.D.E." It matches the as 2nd segment; segments 3-7 distinctly transverse. Pro- original description, and has been labelled "Holotype notum 0.95-0.98x as long as wide, with sides gently female, y e o ius ca i a us Broun, 1881, R.C.Craw det. rounded, subparallel on basal half; disc with diffuse and 1997." indistinct lateral and median sulci, these sometimes obso- lete or obsolescent, and a distinct medio-basal impression Redescription. Length 20.4-24.8 mm, width 8.6-10.8 (Fig. 18). Elytra elongate ovate; intervals 3, 5, and 7 raised mm. as costae; interval 9 raised as costae on apical third and Integument reddish-brown to black; body vestiture of conjoint with interval 3 on declivity. Tarsal segments 1 orbicular to elongate, lineal-lanceolate, white or yellowish
—24— white scales, densest on base of head, sides of pronotum, Paratypes: 2 females, same data as holotype; 3 males, and elytral intervals 2, 4, and 6. Rostrum short and thick, NN, Mt Arthur, 5000 ft [1500 m], 15 December 1961, J.I. 1.5-1.7x apical width in length. Funicle with 1st segment Townsend & G.F. Woods. 1.5-1.8x as long as 2nd segment. Pronotum transverse, 1.1-1.25x wider than long, with sides strongly rounded; Description. Length 20.5-28 mm, width 9.2-12.2 mm. disc with a median basal impression. Elytra elongate ellip- Integument black; body vestiture of pale yellowish- tic; intervals 3, 5, and 7 strongly costate; interval 9 costate white, lineal-lanceolate scales, densest on head and sides on apical quarter; interval 8 flat on basal and apical quar- of pronotum, and forming stripes on elytral disc, declivity, ters, raised medially; intervals 2, 4, and 6 flat, coarsely and sides; secondary scale stripes also present. Rostrum granulate. short and thick, 1.6-1.9x as long as apical width. Funicle Male. Ventrite 5 with a distinct medio-apical impres- with 1st segment 1.6-1.78x as long as 2nd segment (Fig. sion. Median lobe short and stout; aedeagal apodemes 14). Pronotum transverse, 1.15-1.2x wider than long, shorter than median lobe. Endophallus with a distinct, with sides gently to strongly rounded, a median, smooth, inverted V-shaped apical sclerite; basal sclerite without raised area, and a basal impression. Elytra elongate ellip- distinct latero-apical arms (Fig. 53). tic; intervals 3, 5, and 7 strongly costate; secondary costae Female. Ventrite 5 slightly convex basally, with a shal- on intervals 2 and 4 vestigial to strongly developed, and on low impression. Sternite 8 with apodeme short and broad interval 6 absent, vestigial, or strongly developed. (Fig. 71). Hemisternites stout, broad; styli short, broad, Male. Ventrite 5 with an indistinct medio-apical imp- inserted apico-laterally; hemisternal pouches very long ression (Fig. 40). Median lobe stout; aedeagal apodemes (Fig. 78). shorter than median lobe. Endophallus with apical scler- ites; basal sclerite with apico-lateral processes (Fig. 54). Material examined. Holotype, plus 21 non-type exam- Female. Ventrite 5 with an indistinct medio-apical imp- ples (BMNH, CMNZ, LUNZ, NZAC). ression (Fig. 41). Sternite 8 truncate apically; apodeme long. Hemisternites stout, broad; styli short, broad, inser- Distribution. KA, MC, SC, MK: mountain ranges - ted apicolaterally; distal plate with a rod-like proximal Seaward Kaikoura Ra. (Kahutara Saddle), Craigieburn extension; hemisternal pouches long. Ra., Broken R. Basin, Mt Temple, Jollie Peak, Mt Hutt, Mt Somers, Carney's Creek, McCoy's Creek (upper Rangi- Material examined. Type series, plus 158 non-type tata R.), Mt Cook National Park, Mt Dalgety. examples (AMNZ, NZAC). The Kahutara Saddle record is taken from Hunt (1996); no specimens from this area have been examined by the Distribution. NN, BR: mountain ranges - Arthur Ra., author. Specific status of northeastern South Island Domett Ra., Marino Mtns, Braeburn Ra., Matiri Ra., Lyperobius populations require specialist assessment as Glasgow Ra. there may be undescribed taxa in this area. Host plant. Aciphylla ferox (Apiaceae). Host plants. Aciphylla species (Apiaceae). Remarks. Lyperobius clarkei is readily distinguishable Remarks. Although one of the earliest described species, from its sympatric congener L. fallax by size, extensive L. carinatus is very poorly represented in collections. Its development of secondary costae, and genital characters. distributional limits and population status should be ascer- Vicariant populations show consistent geographic vari- tained, as this species - found in the drier and environ- ation in some external characters. Northern populations mentally degraded mid-eastern South Island mountain from Mt Arthur (Arthur Range) south to Mt Owen (Marino ranges — may be endangered. Mountains) are more robust, with strongly rounded pro- notal sides and well developed costae in elytral interval 6. Southern populations (e.g., from the Glasgow Range) are p r b l r ew s ecies slender with gently rounded pronotal sides and the costae in intervals 6 absent to vestigial. The species is named in Figures 5, 11-14,40,41; Map 8 honour of the entomologist C. E. Clarke. Type data. Holotype: female, NN, Mt Arthur, 5000 ft [1500m], 4 February 1965, J.I. Townsend & A.K. Walker, Aciphylla ferox (NZAC), 25.1 x11.8 mm.
—25— p r b x l Kusc e p r b p nd ou Figures 33, 55; Map 9 Figures 15, 19, 56, 72, 79; Map 10 coxalis Kuschel, 1987: 18 (Lyperobius). cupiendus Broun, 1886: 962 (Lyperobius). —Hutton 1904: 203.—Hudson 1923: 390. -Dalla Torre, Schenkling, & Type data. Holotype: male, FD, Hunter Mountains, Mt Marshall 1932: 65. —Hudson 1934: 214. Burns, 1300 m, January 1970, J.I. Townsend (NZAC), aciphyllae Broun, 1917: 442 (Lyperobius), new s n - 19x9.3 mm. n Paratypes (n=121): listed in Kuschel (1987). Type data. Holotype of cupiendus: male, "Mt Tyndall, Redescription. Length 14-23 mm, width 7.3-11 mm. 6500 ft [1950m], GM Thomson" (BMNH), 15.6x9.1 mm. Integument black; body vestiture of greyish-white to Holotype of aciphyllae: female, "Garvie Mts, 7-1- yellowish-white, orbicular to ovate scales, densest on base 1914" (BMNH), 16.8x8.6 mm. of head, elytral intervals 2, 4, and 6, and upper inner face of coxae. Rostrum relatively short and thick, length 1.7- Redescription. Length 12.2-19 mm, width 6.5-9.7 mm. l.8x apical width. Funicle with 1st segment 1.4-1.44× as Integument dark reddish brown to black; body vestiture long as 2nd segment; segments 3-7 transverse. Pronotum of orbicular to ovate, white to pale yellowish-white scales, transverse, 1.19-1.25x wider than long; disc with an ab- densest on base of head, sides and middle of pronotum, breviated median carina or elongate, low, flat, smooth area and elytral intervals 2, 4, and 6. Rostrum relatively short on apical two-thirds, a shallow median basal impression, and thick, length l.6-1.8x apical width (Fig. 15). Funicle and shallow mediolateral impressions. Elytra elongate with 1st segment 2x as long as 2nd segment; 7th segment ovate; intervals 3, 5, and 7 strongly raised as costae; inter- transverse (Fig. 15). Pronotum distinctly transverse, val 9 costate on apical quarter and conjoint with costae on length 0.78- .9× width, with sides strongly rounded (Fig. interval 3; intervals 2, 4, and 6 flat, with sculpture of fine, 19). Elytra ovate; intervals 3, 5, and 7 distinctly raised as low granules. Coxae with a distinct, dense patch of slightly thick, prominent costae; intervals 2, 4, 6, and 8 either flat raised scales on upper half of inner face (Fig. 33). or partially to fully raised as secondary costae; interval 9 Male. Ventrite 5 with a distinct mid-apical impression. costate and conjoint with interval 3 on declivity. Median lobe elongate; aedeagal apodemes distinctly shor- Male. Ventrite 5 with a distinct, shallow, medio-apical ter than median lobe. Endophallus with no apical sclerite; impression. Median lobe stout; aedeagal apodeme longer basal sclerite without apicolateral processes (Fig. 55). than median lobe. Endophallus with apical sclerites; basal Female. Ventrite 5 weakly convex basally, with a shal- sclerite with apicolateral processes (Fig. 56). low medio-apical impression and upturned apical margin. Female. Ventrite 5 with a distinct medio-apical impres- Sternite 8 elongate; apodeme relatively long. Hemisternites sion and upturned apical margin. Sternite 8 with apodeme with stout setae on distal part; hemisternal pouches short. long (Fig. 72). Hemisternites with robust long setae on distal portion; hemisternal pouches short, broad. Bursa Material examined. Type series only (AMNZ, BMNH, copulatrix with accessory lobes (Fig. 79). CMNZ, FRNZ, LUNZ, MONZ, NZAC, OMNZ). Material examined. Holotypes, plus 65 non-type ex- Distribution. FD: mid-eastern to southern ranges from amples (AMNZ, BHPC, BIPB, BMNH, NZAC). west of L. Te Anau to west of L. Poteriteri (Murchison Mtns, Kepler Mtns, Turret Ra., Hunter Mtns, Heath Mtns, Distribution. CO, OL: mountain ranges to west of L. Cleughearn Peak, Kaherekoau Mtns, Cameron Mtns). Wanaka (Cascade Saddle, Mt Aspiring, Mt Tyndall, Headlong Peak, Niger Peak), Pisa Ra., The Remarkables, Host plants. Aciphylla crosby-smithii, A. lyallii, A. pin- Old Woman Ra., Kopuwai / Old Man Ra., Carrick Ra., natifida, Anisotome aromatica, A. capillifolia, A. haastii Garvie Mtns, Um-brella Mtns. (Ap Recorded at 1300 m on the Umbrella Mountains (Dick- inson 1988). Remarks. Lyperobius coxalis is easily distinguished from sympatric congeners by the distinct dense patch of Host plants. Aciphylla congesta, A. lecomtei, A. pinnati- slightly raised scales on the coxae, and from its closest fida, A. simplex, Anisotome flexuosa, A. imbricata (Ap - relatives L. australis and L. nesidiotes by the combination aceae). of elongate -ovate elytra, flat elytral intervals 2, 4, and 6, and relatively short and thick rostrum. — — Remarks. Lyperobius cupiendus is easily distinguished Host plants. Aciphylla congesta,Aciphylla sp. (Apiaceae). from all other species by the combination of small size, relatively thick rostrum, transverse pronotum, funicle with Remarks. Lyperobius eylesi is easily distinguished from first segment twice as long as second segment, and orbi- known allopatric and sympatric congeners by the com- cular to ovate scales on elytral disc. bination of absence of scale patches on the upper inner Elytral fragments (in OMNZ) collected in the face of the coxae, elytral interval 3 strongly costate and Earnscleugh Cave, at the foot of the Old Man Range, conjoint with interval 9 on the declivity, spatulate shape of suggest that this species may have once occurred at lower the female eighth sternite, and male endophallus basal altitudes. sclerite with broad, rounded apicolateral processes. Named for Dr Alan Eyles, who pioneered the use of numerical taxonomy in entomological systematics in New Zealand. y e o ius ey esi ew s ecies Figures 27, 57, 58, 73, 80; Map 11 y e o ius a a ou Type data. Holotype: male, FD, Mt Wilmot, 1350 m, Figures 16, 20, 36, 59; Map 12 Manapouri Exp[edition], Jan[uary] 1970, J. McBurney fallax Broun, 1917: 443 (Lyperobius). -Hudson 1923: (NZAC), 17.9x8.5 mm. 390. —Dalla Torre, Schenkling, & Marshall 1932: 65. Paratypes: 4 females, same data as holotype; 1 male, 4 -Hudson 1934: 214. females, FD, Mt Barber, 1100-1350 m, Manapouri Exp- Type data. Lectotype: female (here designated), MB, [edition], Jan[uary] 1970, J.S. Dugdale. "Mt Miromiro, 6000 ft [1800 m], C.Chilton" (BMNH), Description. Length 16.2-24.9 mm, width 8-11.5 mm. 15.8x8.0 mm. Integument black to dark reddish brown; body vestiture Paralectotype: 1 male (here designated), same data as of pale yellowish-white, oval to elongate-ovate scales, holotype but with no altitude, 15x7.3 mm. densest on head and elytral intervals 2, 4, and 6. Rostrum The female specimen has been designated as lectotype moderately long and thin, 2-2.1 x its apical width. Funicle because the original description states "rostrum with 1st segment 1.6× as long as 2nd segment; segments unimpressed in front of antennae" and "fifth ventral seg- 3-7 rounded. Pronotum transverse, approx. 1.2× as wide ment unimpressed", which are female characters in this as long, with sides broadly rounded and with a distinct, species. shallow, median basal impression. Elytra elongate ovate; intervals 3, 5, and 7 raised as distinct costae; intervals 2, 4, Redescription. Length 13.3-19 mm, width 6.6-9.2 mm. and 6 flat to slightly raised, granulate, transversely rugose; Integument black; body vestiture of pale yellowish to interval 8 flat; intervals 3 and 9 costate and conjoint on white ovate scales, densest on pronotal margins and elytral declivity (Fig. 27). Tarsal segments l and 2 with sole hairs disc and declivity, forming distinct pale stripes on elytra in no longer than those on segment 3, and not greatly pro- intervals 2, 4, 6, and 8 and along margin on apical third, jecting distally. with secondary stripes present on declivity. Rostrum rel- Male. Ventrite 5 with a shallow medio-apical impres- atively thin, 1.95-2.16× apical width in length (Fig. 16). sion. Median lobe short and stout; aedeagal apodemes just Funicle with 1st segment 1.3-1.5× as long as 2nd segment shorter than median lobe. Endophallus with an inverted V- (Fig. 16). Pronotum transverse, 1.1-1.2× wider than long, shaped apical sclerite; basal sclerite with large, rounded with sides strongly rounded (Fig. 20). Elytra elongate apicolateral processes (Fig. 57). ovate; intervals 3, 5, and 7 raised as costae; interval 9 Female. Ventrite 5 convex on basal half, with a distinct costate on apical quarter and conjoint with interval 3. longitudinal impression on apical half; apical margin up- Tarsal segments 1 and 2 with sole hairs not longer than turned. Sternite 8 spatulate, elongate; apodeme relatively those on segment 3, and only slighly projecting distally short (Fig. 73). Hemisternal pouches short. Bursa copu- (Fig. 36). latrix with a sclerite near junction with oviduct (Fig. 80). Male. Ventrite 5 with a distinct mid-apical impression. Median lobe stout; aedeagal apodemes about as long as Material examined. Type series, plus 8 non-type exam- median lobe. Endophallus with apical sclerites (Fig. 59). ples (BHPC, NZAC). Female. Ventrite 5 convex on basal half. Sternite 8 with apodeme short. Hemisternal pouches short. Distribution. FD: Tutoko Vly, Murchison Mtns, Mt Barber, Mt Wilmot, Wilmot Pass (west of L. Manapouri).
—27— Material examined. Type specimens, plus 86 non-type Material examined. Type series only (AMNZ, LUNZ, examples (AMNZ, BMNH, MONZ, NZAC, OMNZ). NZAC).
Distribution. NN, MB, BR, NC: mountain ranges - Distribution. WD: mountains surrounding Fox Glacier Boulder Lake, Tasman Mtns (Domett Ra., Aorere Peak), and Franz Josef Glacier (Mt Moltke, Castle Rocks, Chan- Lockett Ra., Mt Cobb, Peel Ra., Arthur Ra., Owen Ra., cellor Shelf). Marino Mtns, Richmond Ra., Black Birch Ra., St Arnaud Ra., Travers Ra., Victoria Ra., Lewis Pass, Hanmer Ra. Host plant. Aci y a sp. (Apiaceae). (Mt Miromiro), Polar Range (Mt Dome), Arthurs Pass, Broken R. Basin. Remarks. y e o ius g acia is is distinguished by the combination of elytral interval 3 flat on declivity, interval Host plants. Aci y a mo oi A iso ome s (Apiaceae). 7 barely costate and heavily granulate, endophallus basal sclerite without apicolateral processes, and bursa copu- Remarks. y e o ius a a is illustrated in a colour latrix without a sclerite near junction with oviduct. photograph by Meads (1990: 47).
p r b h d n ou p r b l l ew s ecies Figures 6, 17, 21, 25, 37, 61; Map 14 Figures 35, 60; Map 13 u so i Broun, 1914: 127 ( y e o ius —Hudson 1923: 390. —Tillyard 1926: pl. 2 fig. 18. —Dalla Torre, Type data. Holotype: male, WD, Fox Glacier, Chancel- Schenkling, & Marshall 1932: 65. -Hudson 1934: lor Shelf, 1340 m, 13 December 1984, J.W. Early, from 133, pl. XIII fig. 2. base of Aci y a sp. (NZAC), 19.4x8.8 mm. Paratypes: 4 males, same data as holotype, J.W. Early Type data. Lectotype: female (here designated), CO, & P. Syrett; 3 females, same data except 1280 m, 15 Dec "Humboldt Range, 5600 ft [1680 m], GV Hudson" 1984; 1 female, same data except 1280 m, P. Syrett; 1 ( Μ 1 x7 7 female, WD, Westland National Park, Castle Rocks V[ly], 1250 m, 15 Jan 1986, J.W. Early, Aci y a; 1 female, Redescription. Length 11.5-21.5 mm, width 5.9-9.3 mm. WD, Fox Glacier, Chancellor Hut, 1220 m, litter LUNZ Integument black; body vestiture of white or yellowish, 84/8, J.W. Early & P. Syrett; 1 female, WD, Mt Moltke, oval to ovate scales, densest on base of head and sides and 5000 ft [1500m], above Franz Josef Glacier, 20 Jan 1925. middle of pronotal disc, and forming distinct pale stripes in elytral intervals 2, 4, and 6. Rostrum relatively long and Description. Length 18.2-21.9 mm, width 8.1-10.5 mm. thin, 2.06-2.6x its apical width (Fig. 17). Funicle with 1st Integument reddish brown to black; body vestiture of segment twice as long as 2nd segment; segments 3-7 dis- elongate-ovate, yellowish-white scales, densest on head, tinctly transverse, except in Humboldt Mountains popula- sides of pronotum, and elytral intervals 2, 4, and 6. Ros- tions (Fig. 17). Pronotum subsquare, length 0.95-1.07x trum relatively long and thin, 2.2-2.6x its apical width. width, with sides subparallel to gently rounded; disc with Funicle with 1st segment 1.6-1.9× as long as 2nd segment. pale scale stripes in narrow lateral and median longitudinal Pronotum transverse, 1.1-1.3x wider than long, with sides sulci (Fig. 21). Elytra elongate oval to elongate ovate; strongly rounded and with a shallow median basal impres- intervals 3, 5, and 7 raised as costae; interval 9 raised as sion. Elytra elongate elliptic; intervals 3 and 5 distinctly costae on apical quarter; intervals 2, 4, and 6 usually flat, costate on disc; interval 7 flat to slightly raised, strongly sometimes with transverse raised areas joining costae in granulate; interval 3 flat on declivity. intervals 3, 5, and 7 and interrupting the pale stripes. Male. Ventrite 5 with a distinct medio-apical impres- Tarsal segments 1 and 2 with apical sole hairs longer than sion. Median lobe stout; aedeagal apodemes shorter than those on segment 3, and greatly projecting distally (Fig. median lobe. Endophallus with apical sclerites; basal 37). sclerite without apicolateral processes (Fig. 60). Male. Ventrite 5 with a distinct medio-apical impres- Female. Ventrite 5 almost flat medio-apically, with lat- sion. Median lobe stout; aedeagal apodemes about as long eral impressions. Sternite 8 with apodeme relatively long. as median lobe. Endophallus with apical sclerites; basal Hemisternites with distal portion elongate; hemisternal sclerite with apicolateral processes (Fig. 61). pouches more than half length of hemisternites. Female. Ventrite 5 convex on basal half, with a shallow medio-apical impression. Sternite 8 with apodeme long. — — Hemisternites with abundant fine setae on apical portion; Tarsal segments 1 and 2 with sole hairs not much longer hemisternal pouches short and broad. Bursa copulatrix than those on segment 3, only slightly projecting distally. with a distinct sclerotised area near junction with oviduct. Male. Ventrite 5 with a slight medio-apical impression. Median lobe stout; aedeagal apodemes about as long as Material examined. Lectotype, plus 170 non-type ex- median lobe. Endophallus with a pair of narrow apical amples (AMNZ, BHPC, BIPB, BMNH, MONZ, NZAC). sclerites; basal sclerite triangular, with short apicolateral processes (Fig. 62-64). Distribution. CO, OL: mountain ranges - Forbes Mtns Female. Ventrite 5 slightly convex on basal half, with (Mt Tyndall), Humboldt Mtns, Ailsa Mtns, Mt Cardrona, lateral impressions. Tergite 8 broad. Hemisternites broad, Minor Peak, Ben Lomond, Cecil Peak, Eyre Mtns, The stout; hemisternal pouches almost as long as hemisternites. Remarkables, Hector Mtns, Pisa Kopuwai / Old Man Old Woman Garvie Mtns. Material examined. Type series, plus some 200 non- Altitudinal range 1600-1900 m on The Remarkables type examples [approximately one-third of them fragmen- (Patrick et al. 1992), 1600-1690 n the Old Man Range ted] (AMNZ, BHPC, BMNH, CMNZ, LUNZ, MONZ, (Brumley et al. 1986), and 1750-1860 m on the Eyre NZAC, OMNZ). Mountains (Mark et al. 1989). Host plants. Aciphylla dobsonii, A. lecomtei, A. simplex, Distribution. North I.— WN: Wellington, southern coast A. spedenii, Anisotome flexuosa, A. imbricata (Apiaceae). and coastal hills (Happy Valley, Island Bay to mouth of K r r Str n ll S th I — M ΚΑ C Remarks. Elytral pattern and sculpture vary greatly in L. MC, SC: Tarndale, upper Wairau Vly; Black Birch Ra.; hudsoni. In some individuals elytral intervals 2, 4, and 6 Seaward Kaikoura Ra. (Haycock Ra., Mt Fyffe, Kahutara have transverse raised areas joining the costae on the Saddle), Jacks Pass, Hanmer Range; Hanmer Plain; Mt uneven intervals and interrupting the pale scale stripes. Rinser; Craig-ieburn Ra.; Norwell Downs Stn, north of Such individuals predominate in Old Man Range and Burkes Pass, Two Thumb Ra.; Albury Ra.; Grampian Garvie Mountains populations, but are rare in Pisa Range Mtns; Hunters Hills. and Humboldt Mountains populations. This taxon may be The distribution and biogeographic history of this a complex of cryptic species. species were discussed and mapped by Craw (1988).
Host plants. Aciphylla aurea, A. colensoi, A. squarrosa y e o us u o i Pascoe (Apiaceae). Figures 7, 31, 62-64; Map 15 Remarks. A threatened and legally protected species, huttoni Pascoe, 1876: 54 (Lyperobius). -Broun 1880: 445. Lyperobius huttoni is easily recognised by its non-costate -Hutton 1904: 203. -Hudson 1923: 390. —Dalla elytral intervals, sparse body vestiture, and short, thick Torre, Schenkling, & Marshall 1932: 65. —Hudson rostrum. South Island populations, found in montane to 1934: 133. subalpine habitats, have an eastern distribution from the Type data. Lectotype: male (here designated), MB, Black Birch Range (Marlborough) to the Hunters Hills Tarndale ["near the head of the river Wairau, in the Nelson (South Canterbury). province", Captain F.W. Hutton] (BMNH), 20x9 mm. North Island populations were discovered on coastal Paralectotypes (here designated): 2 males, 4 females, cliffs and hills south of Wellington by A.C. O'Connor and same data as lectotype. A.E. Brookes in 1917. These populations once extended Redescription. Length 18-26.2 mm, width 9.2-11.8 mm. from Island Bay to the mouth of the Karori Stream, but by Integument reddish brown to black; body vestiture the late 1980s huttoni was facing extinction in these sparse, of white lineal-lanceolate scales, densest on pro- coastal habitats. A fenced reserve has been established on notal sides and forming pale, reduced stripes only on the south Wellington coast to protect important habitat for elytral declivity. Rostrum relatively short and thick, 1.48- this species (Sherley 1994). Ecology and morphology of 1.69× apical width in length. Funicle with 1st segment Wellington coastal populations have been studied in detail l.33-1.79x as long as 2nd segment; 7th segment trans- (Bull 1967). Hunt (1996) provides further information on verse. Pronotum transverse, with sides broadly rounded; the ecology and conservation status of this species. disc with a mid-basal impression. Elytra elongate elliptic; intervals flat or slightly convex, never distinctly costate.
— 9— p r b nt n ew s ecies of this species can be found in Peat & Patrick (1996) and Figures 8, 28, 34, 65; Map 16 Morris & Hayden (1995).
Type data. Holotype: male, FD, Gertrude Saddle, 1350 p r b n d t Kusc e 3 n r 19 1 tr ( AC 1 3x 1 Paratypes: 1 male, FD, Homer Tunnel, 100 m, 5 Dec Figure 66; Map 17 1987, B.H. Patrick; 1 male, 6 females, FD, Fiordland N.P., esi io es Kuschel, 1987: 20 ( y e o ius Gertrude Vly, 1150-1380 m, 6 Feb 1980, J.W. Early, R. M. Emberson, & P.T. Townsend; 4 males, 5 females, FD, Type data. Holotype: male, The Snares, Broughton I., 14 Gertrude Saddle, Homer, 4500 ft [1350 m], 5 Feb 1963, January 1977, D.S. Horning (NZAC), 21.0x10.0 mm. J.I. Townsend, Aci y a co ges a & A iso ome aas ii; 1 Paratypes ( =11 listed in Kuschel (1987). male, FD, Gertrude Saddle, Homer Tunnel, 4830 ft [1450 Redescription. Length 20.5-22.5 mm, width 9.3-10.0 m], 5 Feb 1963, R.M. Bull. Description. Length 13.4-21.8 mm, width 6.3-10.5 mm. Integument black, with elytra and tarsi often dark red- Integument black to dark reddish brown; body vestiture dish brown; body vestiture of orbicular to ovate, greyish- of ovate, pale yellowish-white scales, densest on head, white to yellowish-brown scales, densest on elytral inter- sides of pronotum, and intervals 2, 4, and 6, but nowhere vals 2, 4, and 6, elytral declivity, and upper inner face of as abundant as in most other species. Rostrum relatively coxae. Rostrum relatively long and thin, 2.25-2.42x its long and thin, 2.0-2.3× its apical width. Funicle with 1st apical width. Funicle with 1st segment 1.58 —1.76× as long seg-ment 1.8-2.2x as long as 2nd segment. Pronotum as 2nd segment; segments 3-7 becoming progressively about as long as wide, with sides gently rounded. Elytra more transverse. Pronotum slightly transverse, l.1-1.19x elongate ovate; intervals 3, 5, and 7 weakly raised as low wider than long; disc with a median carina or elongate, flat, costae; intervals 3 and 9 flat on apical third (Fig. 28); smooth line often extending length of pronotum, and with intervals 2, 4, and 6 very slightly raised and transversely a median basal impression. Elytra elongate elliptic; inter- rugose. Tarsal segments 1 and 2 with apical sole hairs vals 3, 5, and 7 raised as low costae; interval 3 with costae longer than those of segment 3, and greatly projecting obsolete to obsolescent on apical third; interval 9 flat on distally. apical quarter; intervals 2, 4, and 6 flat, convex or sub- Male. Ventrite 5 with a distinct medio-apical impres- costate. sion. Median lobe stout; aedeagal apodemes about as long Male. Ventrite 5 with a very light medio-apical impres- as median lobe. Endophallus with apical sclerites; basal sion. Median lobe robust, elongate; aedeagal apodemes sclerite with short apicolateral processes (Fig. 65). amost as long as median lobe. Endophallus without apical Female. Ventrite 5 weakly convex on basal half, at most sclerites; basal sclerite without apicolateral processes shallowly impressed on apical half. Sternite 8 with apo- (Fig. 66). deme relatively short. Hemisternal pouch short, narrow. Female. Ventrite 5 strongly convex basally, with a deep, Bursa copulatrix with a distinct sclerotised area near transverse medio-apical impression and upturned apical junction with oviduct. margin. Sternite 8 with apodeme relatively long. Hemi- sternites with stout setae on proximal part; hemisternal Material examined. Type series, plus 88 non-type ex- pouches short, broad. amples (AMNZ, BHPC, LUNZ, MONZ, NZAC). Material examined. Type series only (CMNZ, LUNZ, Distribution. WD, FD: mountain ranges — Olivine Ra., MONZ, NZAC). Red Hill Ra., Darran Mtns, Gertrude Saddle, Homer Sad- dle, Mackinnon Pass, Murchison Mtns. Distribution. The Snares (Broughton I. only).
Host plants. Aci y a co ges a A iso ome aas ii (Api- Host plant. A iso ome acu i o ia (Apiaceae). aceae). Remarks. The rarest of all y e o ius species, esi i- Remarks. y e o ius mo a us is easily distinguished o es is confined to a small islet in The Snares, where it is from all other species by the combination of elytral in- found in a patch of host plant not much larger than a small tervals 3 and 9 flat on apical quarters (Fig. 28) and much room (J.W. Early, pers. comm.). less abundant pale scaling on elytra. Colour photographs —3 --3 — p r b p tr ew s ecies elongate habitus, lineal-ovate to lineal-lanceolate hairlike Figures 22, 42, 43, 67, 74, 81; Map 18 scales, and non-transverse third tarsal segment (in male), and from its only known sympatric congener L. barbarae Type data. Holotype: male, CO, Kakanui Mountains, by size, pronotal sculpture and pattern, and secondary Otepopo Spur, 1240 m, 13 December 1989, B.H. Patrick, l tr p n v n l tr l nt rv l Α l r ph t r ph on A[ciphylla] gracilis (NZAC), 21.7x10.3 mm. of this species can be found in Patrick (1991, fig. 9). Paratypes: 2 males, 1 female, CO, St Mary's Ra., Awa- Named for entomologist Brian Patrick, who has con- kino Ski Field, 1220 m, 18 Feb 1989, A.C. Harris, Aci- tributed new distribution and host plant records plus num- phylla aurea; 1 female, CO, St Mary's Ra., 19 Feb 1989, erous specimens towards this revision. B.H. Patrick; 5 females, CO, Little Mt Domet, 1150-1600 m, 21 Mar 1991, B.H. P[atrick], feeding on A[ciphylla] aurea. p r b p d n ou Figures 9, 23, 68, 75, 82; Map 19 Description. Length 22.3-26.5 mm, width 10-12.5 mm. Integument dark reddish brown; body vestiture of yel- spedenii Broun, 1917: 443 (Lyperobius). —Hudson 1923: lowish-grey, lineal-ovate to lineal-lanceolate scales (often 390. —Dalla Torre, Schenkling, & Marshall 1932: 65. hairlike), densest on head and elytral intervals 2, 4, and 6. —Hudson 1934: 214. Rostrum relatively short and thick, length 1.7-1.95x Type data. Holotype: female, SL, "Garvie Mts., nr. Blue apical width. Funicle with 1st segment 1.33-1.6x as long Lake, J. Speden, 7.1.1914" (BMNH), 25.2x12.1 mm. as 2nd segment; segments 3-6 transverse; segment 7 roun- Redescription. Length 17.7-23.8 mm, width 9.1-11.3 ded. Pronotum weakly transverse, l.07-1.19x wider than long, broadly rounded, with an abbreviated median carina Integument black; body vestiture of orbicular to ovate, (Fig. 22). Elytra elongate elliptic; intervals 3, 5, and 7 pale yellowish-white to grey yellowish-white scales, den- raised as costae; intervals 2, 4, and 6 with large granules, sest on head and even elytral intervals. Rostrum variable these sometimes confluent and forming vestigial to secon- in length, 1.76-2.5× its apical width. Funicle with 1st dary costae. segment 1.5-1.6x as long as 2nd segment; segments 3-6 Male. Third tarsal segment about as long as broad. Ven- transverse; segment 7 variably transverse to about as long trite 5 with a deep, circular, medio-apical impression (Fig. as wide. Pronotum transverse, 1.1-1.24x wider than long, 42). Median lobe short, stout; aedeagal apodemes about as with sides gently rounded and then subparallel to sinuate; long as median lobe. Endophallus without apical sclerites; disc with pale scales scattered in no particular pattern, an basal sclerite with apicolateral processes (Fig. 67). elongate, low, flat, smooth median strip, and a distinct Female. Ventrite 5 with a deep, circular, medio-apical median basal impression (Fig. 23). Elytra broadly ovate; impression and upturned apical margin (Fig. 43). Sternite intervals 3, 5, and 7 costate; even intervals with large gran- 8 stout; apodeme short and thick (Fig. 74). Hemisternites ules, these sometimes confluent and forming secondary to with stout setae on proximal portion; hemisternal pouches vestigial costae; secondary scale stripes also present. short, broad. Bursa copulatrix with accessory lobes (Fig. Male. Third tarsal segment transverse. Ventrite 5 with 81). a medio-apical, obconical to semicircular impression. Median lobe very stout; aedeagal apodemes about as long Material examined. Type series, plus 16 non-type ex- as median lobe. Endophallus without apical sclerite(s); amples (BHPC, BIPB, NZAC, OMNZ). basal sclerite with apicolateral processes (Fig. 68). Female. Sternite 8 elongate; apodeme relatively long Distribution. SC/MK, CO, DN: mountain ranges - and thin (Fig. 75). Hemisternites with stout setae on apical Grampian Ra., St Mary's Ra., Little Mt Domet, Ida Ra., portions; hemisternal pouches short. Bursa copulatrix Kakanui Ra., Pig Root Creek. with accessory lobes (Fig. 82).
Host plants. Aci y a au ea Α mo a a var. gracilis Material examined. Holotype, plus 36 non-type examples (Apiaceae). (BHPC, BIPB, BMNH, NZAC).
Remarks. First collected in 1964, Lyperobius patricki Distribution. MC, MK, WD, CO, OL, SL: mountain occurs as low as 800 m in North Otago and South Can- ranges — Mt Hutt Ra., Godley Glacier, Young Ra. (Mt terbury ranges. This large species is distinguished from its Brewster), Mark Ra., ranges around L. Ohau, Forbes Mtns vicariant relative spedenii by its larger and more (Headlong Peak), upper Dart R., Harris Mtns (End Peak,
—31— Niger Peak), The Remarkables, Eyre Mtns, Snowdon Material examined. Type series only (BHPC, NZAC). Peak, Garvie Mtns, Takitimu Mtns. Altitudinal range 1700-1800 m on The Remarkables Distribution. SL: Takitimu Mtns (Family Peak, Mt (Patrick e a 1992), 1400-1860 m on the Eyre Mountains Brown, Spence Peak). (Mark e a 1989), and 1250-1420 m on the Takitimu M nt n ( tr p r Host plants. A iso ome aas ii A i i e a (Apiaceae).
Host plants. Aci y a ecom ei A i a i i a A s e - Remarks. The elegant and elongate habitus (Fig. 10) and e ii A sim e (Apiaceae). the genitalia of ow se i serve to distinguish this dis- tinctive species immediately from all others in the genus. Remarks. y e o ius s e e ii is the most widely dis- Named for Mr J.I. Townsend, who collected many of tributed of the southern species. A colour photograph the specimens upon which this revision is based. appears in Peat (1991).
y e o ius ow se i ew s ecies E E E CES Figures 10, 24, 69, 76, 83; Map 20 Alonso-Zarazaga, M.A. 1988: Caracteres differenciales de los generos o y ius Dejean, 1821 y C eo i us Type data. Holotype: female, SL, Takitimu Mountains, Schoenherr, 1823 con descripcion de un nuevo repres- 4000 ft [1200 m], 12 February 1963, A iso ome J.I. entante i e ico C eo i us se a oi n. sp. (Coleoptera: Townsend (NZAC), 18.6x8 mm. Curculionidae: Brachyderinae). Paratypes: 3 males, same data as holotype; 1 male, SL, Gio a e I a ia o i E omo ogia 123-129. Takitimu [Mtns], 1500 m, 24 Jan 1981, B.H. Patrick; 2 males, 2 females, type locality, 4300 ft [1290 m], 30 Nov Anderson, R.S. 1993: Weevils and plants: phylogenetic 1962, J.I. Townsend & A.C. Eyles, A iso ome; 2 males, 2 versus ecological mediation of evolution of host plant females, type locality, 5000 ft [1500 m], 12 Feb 1963, associations in Curculioninae (Coleoptera: Curculion- R.M. B[ull]. idae). Memoi s o e E omo ogica Socie y o Ca - a a 1 5 197-232. Description. Length 13.6-19.8, width 6.1-8.4 mm. Armstrong, J.F. 1870: On the vegetation of the neigh- Integument black or dark reddish brown; body vestiture bourhood of Christchurch, including Riccarton, Dry of oval to ovate white scales, densest on sides, middle of Bush, etc. a sac io s a ocee i gs o e ew pronotal disc, and elytral intervals 2, 4, and 6. Rostrum ea a I s i u e 2: 119-128. relatively long and thin, 2.21-2.58x its apical width. Baker, J.; Seward, D. 1996: Timing of Cretaceous exten- Funicle with 1st segment 1.8-2.0x as long as 2nd segment. sion and Miocene compression in northeast South Pronotum 0.94-1.08× as long as wide, with sides gently Island: constraints from Rb-Sr and fission-track dating rounded; disc with 3 longitudinal scale stripes lying in of an igneous pluton. ec o ics 15 976-983. distinct shallow sulci (Fig. 24). Elytra elongate elliptic; intervals 3, 5, and 7 raised as distinct costae; interval 9 Barratt, B.I.P. 1985: Coleoptera. I Entomological sur- raised as a distinct costa on apical quarter and conjoint vey of the Garvie Mountains. Dunedin, Otago Section, with interval 3 on declivity; intervals 2, 4, and 6 flat, never Entomological Society of New Zealand. with raised transverse areas. Tarsal segments 1 and 2 with Baum, D.A.; Donoghue, M.J. 1995: Choosing among alt- sole hairs longer than those on segment 3, and greatly pro- ernative "phylogenetic" species concepts. Sys ema ic jecting distally. o a y 560-573. Male. Ventrite 5 with a distinct medio-apical impres- Bennett, B.G. 1987: Observations on rearing Ka oco e s sion. Median lobe elongate; aedeagal apodemes longer i os o i (Coleoptera: Curculionidae: Molytinae). than median lobe. Endophallus with apical sclerites; basal ew ea a e omo ogis 9: 34-37. sclerite with short apical processes (Fig. 69). Berenbaum, M.R. 1990: Evolution of specialization in Female. Ventrite 5 with a shallow medio-apical impres- insect-umbellifer associations. A ua e iew o e o- sion. Sternite 8 elongate, with apodeme long (Fig. 76). mo ogy 35 319-343. Hemisternal pouch short and broad. Bursa copulatrix with a distinct sclerotised area near junction with oviduct (Fig. Broun, T. 1880: Manual of the New Zealand Coleoptera 83). [Part l]. Wellington, Colonial Museum and Geologi- cal Survey Department. xix + 651 pp. —32--3 — — 1881: Manual of the New Zealand Coleoptera, Part Dalla Torre, K.W. von; Schenkling, S.; Marshall, G.A.K., 2. Wellington, Colonial Museum and Geological Sur- Sir 1932: Curculionidae: Subfam. Hylobiinae. Co eo- vey Department. Pp. i-xxiii, 653-744. e o um ca a ogus 1 —1886: Manual of the New Zealand Coleoptera, Parts Darwin, C. 1859: On the origin of species by means of 3 & 4. Wellington, Colonial Museum and Geological natural selection, or the preservation of favoured races Survey Department. Pp. i-xvii, 745-973. in the struggle for life. London, John Murray. 1911: Additions to the coleopterous fauna of the Dawson, J.W. 1971: Relationships of the New Zealand Chatham Islands. a sac io s a ocee i gs o Umbelliferae. o a ica ou a o e i aea Soci- e ew ea a I s i u e 3 92-115. e y (su eme 1 43-62. — 1914: Descriptions of new genera and species of Dickinson, K.J.M. 1988: Umbrella Ecological District. Coleoptera. u e i o e ew ea a I s i u e ew ea a o ec e a u a A eas og amme 1( 79-142. o 7. Wellington, Department of Conservation. 1917: Descriptions of new genera and species of Dugdale, J.S. 1975: The insects in relation to plants. Pp. Coleoptera. u e i o e ew ea a I s i u e 561-589 i G. Kuschel (ed.), Biogeography and eco- 1(5 347-474. logy in New Zealand. The Hague, Dr W. Junk. Brown,J.H. 1995: Macroecology. 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Hunt, M.R. 1996: The distribution and habitat require- ments of the large speargrass weevil ( y e o ius 1987a: Immature stages of Curculionidae (Coleo- u o i and characteristics of host speargrass plants ptera): the larva and pupa of Ka oco e s i os o i (Aci y a squa osa and their vegetation associations. (Molytinae). ew ea a e omo ogis 9: 29-34. Unpubl. MSc thesis, Victoria University of Wellington. — 1987b: Immature stages of Curculionoidea (Col- Hutton, F.W. 1904: Index faunae Novae Zelandiae. Lon- eoptera): rearing records 1964-1986. ew ea a don, Dulau & Co. e omo ogis 9 44-56. Kluge, A.G.; Farris, J.S. 1969: Quantitative phyletics and — 1993: Larvae of Curculionoidea (Insecta: Coleo- the evolution of anurans. Sys ema ic oo ogy 1 1- ptera): a systematic overview. au a o ew ea a 32. o Kuschel, G. 1971: Entomology of the Aucklands and other Meads, M. 1990: Forgotten fauna: the rare, endangered, islands south of New Zealand: Coleoptera : Curcu- and protected invertebrates of New Zealand. Welling- lionidae. aci ic i sec s mo og a 27: 225-259. ton, DSIR Publishing. 1987: The subfamily Molytinae (Coleoptera: Curcu- — 1994: Translocation of New Zealand's endangered lionidae): general notes and descriptions of new taxa insects as a tool for conservation. Pp. 53-56 i M. from New Zealand and Chile. ew ea a e omo o- Serena (ed.), Reintroduction biology of Australian gis 9 11-29. and New Zealand fauna. Chipping Norton, N.S.W., Surrey Beatty & Sons. Lacordaire, T. 1863: Histoire naturelle des insectes. Gen- era des Coleoptères. Vol. 6. Paris, Roret. Metcalfe, C.R.; Chalk, L. 1950: Anatomy of the dicoty- ledons. Vol. 1. Oxford, Clarendon Press.
-3 - Morimoto, K. 1962: Key to families, subfamilies, tribes Ramsay, G.W.; Meads, M.J.; Sherley, G.H.; Gibbs, G.W. and genera of the superfamily Curculionoidea of Japan 1988: Research on terrestrial insects of New Zealand. excluding Scolytidae, Platypodidae and Cossoninae. W G esea c e iew o 1 Wildlife Research ou a o e acu y o Ag icu u e Kyus u U i- Liaison Group. e si y 1 21-66. Rance, B.D.; Patrick, B.H. 1988: Botany and Lepidoptera 1982: The family Curculionidae of Japan. 1. Sub- of The Snares. Invercargill, Department of Conser- family Hylobiinae. Esakia o 19 51-121. vation. Morris, R.; Hayden, P. 1995: Wild South's living treas- Roberts, W.H. [?Sherwood] 1895: Southland in 1856-57 ures of New Zealand. Auckland, HarperCollins. with a journey from Nelson to Southland in 1856. Morrone, J.J.; Crisci, J.V. 1995: Historical biogeography: Invercargill, Southland Times. introduction to methods. A ua e iew o eco ogy Sanderson, M.J. 1989: Confidence limits on phylogenies: a sys ema ics 26: 373-401. the bootstrap revisited. C a is ics 5 113-129. Nelson, G. 1973: Classification as an expression of phy- Schenkling, S.; Marshall, G.A.K. 1931: Curculionidae: logenetic relationships. Sys ema ic oo ogy 344- Cylindrorrhininae. Pp. 1-23 i Coleopterorum cata- 359. logus, pars 114. Berlin, W. Junk. rr ; rnb ll I Μ 1993 C n z b n dj - Scherf, H. 1964: Die Entwicklungsstadien der Mitteleuro- nt t n v lv n pl t b nd r th t päischen Curculioniden (Morphologie, Bionomie, l nd p 51- 7 i P.F. Ballance (ed.), South Ökologie). A a u ge e Se cke e gisc e Pacific - Sedimentary basins of the world, vol. 2. a u o sc e e Gese sc a 5 Amsterdam, Elsevier. Schoenherr, C.J. 1823: Tabula synoptica familiae curculi- Pascoe, F.P. 1876: Descriptions of new genera and species onidum. Pp. 1132-1146 i Isis von Oken. of New Zealand Coleoptera (Part II). A a s a mag- Schöps, K.; Emberson, R.M.; Wratten, S.D. 1998: Does a i e o a u a is o y ( 17 48-60. host-plant exploitation influence the population — 1877: Descriptions of new genera and species of dynamics of a rare weevil? Pp. 119-123 i New Zealand Coleoptera (Part IV). A a s a mag- Baumgartner; Brandmyr, P.; Manly, B.F.J. (eds), a i e o a u a is o y ( 19 140-147. Population and community ecology for insect Patrick, B.H. 1991: Insects of the Dansey Ecological Dis- management and conservation. A.A. Balkerma, trict. Scie ce a esea c se ies o 3 Wellington, Rotterdam. Department of Conservation. Sherley, G. 1994: Threatened insect conservation: the his- —1994: Hawkdun Ecological District invertebrate sur- torical perspective. oC ea e e s ecies occasio a vey. Scie ce a esea c se ies o Wellington, u ica io o 2-4. Department of Conservation. Smreczynski, S. 1968: Coleoptera: Curculionidae. K uc e Patrick, B.H.; Lyford, B.M.; Ward, J.B.; Barratt, B.I.P. o O ac a ia Owa ow o ski C esc I (9 c 1992: Lepidoptera and other insects of the Rastus Solari, F. 1941: Revisione dei eo i us Italian ed Burn Basin, The Remarkables, Otago. ou a o e alcune note di sistematica generale dei Curculionidi. oya Socie y o ew ea a 22: 265-278. Memo ie della Socie a E omo ogica I a ia a Patrick, B.H.; Rance, B.D.; Lyford, B.M.; Barratt, B.I.P. 43-90. 1987: Entomological survey of Snowdon Peak - Liv- Stiassny, M.L.J.; de Pinna, M.C.C. 1994: Basal taxa and ingstone Ecological District, New Zealand. Unpubl. the role of cladistic patterns in the evaluation of con- report to Department of Conservation. servation priorities: a view from fresh water. Pp. 235- Peat, N. 1991: Protecting an icon. o es a i ( 249 i P.L. Forey, C.J. Humphries, & R.I. Vane- 36-41. Wright (eds), Systematics and conservation evaluation. Oxford, Clarendon Press, Peat, N.; Patrick, B.H. 1996: Wild Fiordland. Dunedin, University of Otago Press. Styles, J.H. 1973: Notes on New Zealand weevils. ew ea a e omo ogis 5 351-359. Penny, D.; O'Kelly, C.J. 1991: Seeds of a universal tree. a u e 35 106-107. Suggate, R.P.; Stevens, G.R.; Te Punga, M.T. (eds) 1978: The geology of New Zealand (2 vols). Wellington, Government Printer.
-35- Sutherland, R. 1996: Transpressional development of the Worthy, T.H. 1997: Quaternary fossil fauna of South Can- Australia-Pacific boundary through southern South terbury, South Island, New Zealand. ou a of e Island, New Zealand: constraints from Miocene-Plio- oya Socie y of ew ea a 7 67-162. cene sediments, Waiho-1 borehole, South Westland. ew ea a ou a of geo ogy a geo ysics 39 251-264. Swofford, D.L. 1993: PAUP: Phylogenetic Analysis Using Parsimony, version 3.4. Champaign, Illinois Natural History Survey. Thomas, B.W. 1996: Working with weevils. ew ea a scie ce o y o em e 199 1 -11 Thomas, B.W.; Meads, M.J.; Notman, P.R. 1992: A report on the restoration of knobbled weevils ( a am us s i oca ae and flax weevils (A ago us ai u i to Breaksea Island, Breaksea Sound, Fiordland. SI a esou ces ec ica eco 79 Tillyard, R.J. 1926: The insects of Australia and New Zealand. Sydney, Angus & Robertson. Tisdall, C. 1994: Setting priorities for the conservation of New Zealand's threatened plants and animals. Wel- lington, Department of Conservation. Travers, W.T.L. 1869: Remarks on a comparison of the general features of the flora of the provinces of Nelson and Marlborough with that of Canterbury. a sac io s of e ew ea a I s i u e 1 ( e 174-179. Vaurie, P. 1973: The weevil genera oma i o us and O o e us of the Neotropical Cholinae (Coleoptera, Curculionidae). u e i of e Ame ica Museum of a u a is o y 15 (1 1 -49. Wardle, P. 1968: Evidence for an indigenous pre-Quater- nary element in the mountain flora of New Zealand. ew ea a ou a of o a y 120-125. 1978: Origin of the New Zealand mountain flora, with special reference to trans-Tasman relationships. ew ea a ou a of o a y 1 535-550. Watrous, L.E.; Wheeler, Q.D. 1981: The out-group com- parison method of character analysis. Sys ema ic oo- ogy 3 1-11 Wells, S.M.; Pyle, R.M.; Collins, M.N. 1983: The IUCN invertebrate red data book. Mauges, IUCN. Williams, P.H.; Humphries, C.J.; Vane-Wright, R.I. 1991: Measuring biodiversity: taxonomic relatedness for conservation priorities. Aus a ia sys ema ic o a y 665-679. Williams, P.H.; Vane-Wright, R.I.; Humphries, C.J. 1993: Measuring biodiversity for choosing conservation areas. Pp. 309-328 i J. LaSalle & I. Gauld (eds), Hymenoptera and biodiversity. Wallingford, CAB International. —36— A E I 1 Off h r M l t n nd r l t d t x x n d n r l t n t ph l n t n l
Ac ees gy e a i Pascoe Philippine Islands i us Herbst Europe A. i ayami (Kono) East Asia ge ma us (Linnaeus) Europe Ac ees sp. Vanuatu g a io os is Kuster Europe A iso y c us a a a us Rossi Europe se ia o u c a us Heyd. Turkestan A. mo ac us Germar Europe L. tenebrionoides Pallas Europe A e y o ius asa is (Heller) Borneo, Mt Kinabalu Marshallius bondari Rosado- t Brazil A. ci c us Marshall Borneo, Mt Kinabalu M ic u a us (Germar) Brazil A i icus g a ia o Pascoe Brazil Me a y o ius u igi osus A y o io es (Boheman) Paraguay Morimoto Japan y es i e sus (Pascoe) Brazil O o i us ae io s Boisduval Australia B. g ammicus (Pascoe) Peru O c ucia us Montrouzier New Caledonia i u us (Pascoe) [no locality] O cy i i os is (Fabricius) Australia o i e (Boheman) Brazil O g a os a sus Fairmaire Fiji e usus (Boheman) Brazil O k ugi Boheman Tasmania sc mi ii (Boheman) Brazil ac y o ius ici o us (Germar) N. America a a (Pascoe) Venezuela agio oeus o ie a is Ca e ius u e osus (Fairmaire (Motschulsky) Formosa & Germain) Chile a ameco s a i osa Wiedman India Ce o o osus Schoenherr Brazil s a e iae (Marshall) S. Africa ysce us e o ga us (Roelofs) Japan isso es iceae Illiger Europe ae i e is (Hustache) Japan e es a o i ea a Pascoe Colombia o ie a is (Motschulsky) Japan a e a s (Guérin) Colombia D. e o a us (Roelofs) Japan P. sca osus (Pascoe) Colombia E is o us umi us Kirsch Columbia u guicu a us (Boheman) Colombia Eu ocimi us ma e eimii P. e ucosus Pascoe Colombia (Boheman) N. America o o y o ius eae Faust Burma Eu ycus a o icus Heller Japan ecas s u sus Champion Guatemala E e e u yi Marshall Malay Peninsula Sy o us u gi us Pascoe French Guiana ei i o us e ico is (Hustache) Brazil, Argentina c oicus (Germar) Argentina e e a gi (Hustache) Argentina e y o us (Klug) [no locality] KEY O GE E A A S ECIES O MO Y I I i ica us (Boheman) Argentina K OW OM EW EA A o yc i us (Germar) Paraguay ei i us ac a ius Germar Paraguay 1 Prothorax and elytra with tubercles and knobs (Fig. 1- esyc o ius e u osus Marshall Western Samoa 4); ocular lobes well developed and partly covering esyc o ius sp. Vanuatu eyes; metasternum in front of hind coxae with a dis- ossi Chujo Japan tinct projecting tooth (Fig. 29) ... Hadramphus .. 2 y o i e us ge e i (Boheman) Japan —Prothorax and elytra without tubercles and knobs (Fig. a o i (Faust) Japan 5-10); ocular lobes weakly developed and not partly mo a us (Kono) Japan covering eyes; metasternum in front of hind coxae y o ius a ie is (Linnaeus) Europe without a distinct projecting tooth but often with a co ge e (Dalla Torre, slight to prominent overhang (Fig. 30, 31) Schenkling & Marshall) N. America Lyperobius .. 5 a es (Herbst) N. America 2(1) Elytra (in dorsal view) straight sided, widest at apex; Isc ioma us cos i e is Hustache Colombia 2nd segment of funicle about as long as 1st segment I e u a s Faust Colombia ... (p. 21) .. H. pittospori I i oi es Kirsch Ecuador —Elytra (in dorsal view) rounded on sides, widest near i a us a e sis Reitter Europe middle; 2nd segment of funicle distinctly shorter than co o a us Goe e Europe 1st segment ... 3
—37— 3(2) Elytra with no tubercle on interval 3 in declivity, and 13(12) Pronotal disc with raised areas (if present) largely no tubercle on interval 9 opposite articulation of vent- confined to a median carina or low, flat, narrow ele- rites 4 and 5 ... (p. 22) .. H. tuberculatus vation that may be abbreviated, and with pale scales —Elytra with a tubercle on interval 3 in declivity and a scattered in no particular pattern; pronotal sides (in tubercle on interval 9, just above articulation of vent- dorsal view) subparallel to sinuate (Fig. 22, 23) ... 14 rites 4 and 5 .4 —Pronotal disc with a distinct, basal to medio-apical, broadly triangular area of low, raised, sparsely punc- 4(3) Elytral interval 7 with spinose tubercles; pronotum tate elevations that may be separated by longitudinal with irregular flattened granules; elytra without gran- sulci on either side of a median carina, and with pale ules ... (p. 21) .. H. spinipennis scales forming diffuse to distinct, broad lateral stripes; —Elytral interval 7 with conical tubercles; pronotum and pronotal sides (in dorsal view) narrowly to broadly elytra with conspicuous shiny raised granules rounded (e.g., Fig. 20) ... 15 ... (p. 22) .. H. stilbocarpae 14(13) Elytra elongate elliptic, with disc scales elongate 5(l) Elytra without any intervals raised as distinct costae ovate to linear lanceolate; male 3rd tarsal segment (Fig. 7) ... (p. 29) .. L. huttoni almost as long as broad ... (p. 31) .. L. patricki —Elytra with at least intervals 3 and 5 raised as distinct —Elytra elongate ovate, with disc scales oval to ovate; costae (e.g., Fig. 5, 6, 8, 9) male 3rd tarsal segment (in dorsal view) much broader 6(5) Coxae with a distinct dense patch of slightly raised than long ... (p. 31) .. L. spedenii scales on upper half of inner face (Fig. 33) ... 7 15(14) Elytra elongate ovate; rostrum relatively long and —Coxae without a distinct dense patch of slightly raised thin (Fig. 16); hind tibiae distinctly sinuate; small scales on upper half of inner face (Fig. 34) ... 9 species, 14-19 mm in length ... (p. 27) .. L. fallax 7(6) Elytral intervals 3 and 9 (on declivity) flat or very —Elytra elongate elliptic; rostrum short and thick (Fig. slightly raised ... (p. 30) .. L. nesidiotes 14); hind tibiae not distinctly sinuate; large species, —Elytral intervals 3 and 9 (on declivity) raised as distinct 20-28 mm in length ... 16 conjoint costae ... 8 16(15) Ventrite 5 in male with a distinct medio-apical im- 8(7) Pronotum with sides broadly rounded; elytra elon- pression, in female with basal half distinctly convex gate elliptic, with even and uneven intervals raised as (in lateral view) ... (p. 24) .. L. carinatus costae ... (p. 23) .. L. australis —Ventrite 5 in male with an indistinct medio-apical im- —Pronotum with sides subparallel; elytra broadly elon- pression (Fig. 40), in female with basal half plane (in gate ovate, with only uneven intervals on disc raised as lateral view) ... (p. 25) .. L. clarkei costae ... (p. 26) .. L. coxalis 17(11) Rostrum short and thick (length 1.6-1.7x width) 9(6) Elytral declivity and apex with all intervals almost (Fig. 15); elytra oval; pronotum much wider than long flat to slightly convex (Fig. 28) ... 10 (Fig. 19) ... (p. 26) .. L. cupiendus —Elytral declivity and apex with at least intervals 3 and —Rostrum moderately elongate and narrow (length 5, and often 9 as well, distinctly costate (Fig. 27) ... 11 greater than 2x width) (Fig. 17); elytra elongate ovate to elongate elliptic; pronotum about as long as wide or 10(9) Elytral interval 7 granulate... (p. 28) .. L. glacialis slightly longer than wide ... 18 —Elytral interval 7 smooth ... (p. 30) .. L. montanus 18(17) Elytra elongate elliptic; pronotal disc with lateral 11(9) Elytral disc with secondary scale stripes between sulci interrupted by raised areas just before apical con- striae and costae; main scale stripe composed of scat- striction (Fig. 24) ... (p. 32) .. L. townsendi tered pale scales (Fig.26) ... 12 —Elytra elongate ovate; pronotal disc with lateral sulci —Elytral disc without secondary scale stripes between and pale scale stripes not interrupted by raised areas striae and the costae; main scale stripes composed of just before apical constriction (Fig. 18, 21) ... 19 densely packed pale scales (Fig. 25) ... 17 19(18) Pronotal disc with lateral and median sulci diffuse 12(11) Even elytral intervals with transverse raised areas and indistinct, sometimes obsolete or obsolescent (p. 27) .. L. eylesi (Fig. 18) ... (p. 24) .. L. barbarae —Even elytral intervals with rows of large granules that —Pronotal disc with lateral and median sulci narrow and sometimes are coalesced to form vestigial to second- distinct (Fig. 21) ... (p. 28) .. L. hudsoni ary costae, especially on basal half of disc ... 13
-3 - I US A IO S
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— 6— O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ `` ee Ki gs ❑ C a ams ❑ C a ams ■ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ A l nd ❑ A l nd ❑ Cam e ❑ Cam e ❑
Ma 1 Co ec io oca i ies a am us i os o i. Ma Co ec io oca i ies a am us s i i e is.
O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ■ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ A l nd ❑ A l nd ❑ Cam e ❑ Cam e ❑
Ma 3 Co ec io oca i ies a am us s i oca ae. Ma Co ec io oca i ies a am us u e cu a us.
—57—
O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑
ee Ki gs ❑ ` ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y Li A i o es i A i o es ❑ A l nd ❑ A l nd ❑ Cam e ❑ Cam e ❑
Ma 5 Co ec io oca i ies y e o ius aus a is. Ma Co ec io oca i ies y e o ius a a ae.
O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ A l nd ❑ A l nd ❑ ❑ Cam e ❑ Cam e
Ma 7 Co ec io oca i ies y e o ius ca i a us. Ma Co ec io oca i ies y e o ius c a kei.
-5 - O S O E O S O E IS A S IS A S
Ke ma ecs ❑ Ke ma ecs ❑
ee Ki gs ❑ ee Ki gs ❑
C a ams ❑ C a ams ❑
S a es ❑ S a es ❑
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A i o es ❑ A i o es ❑
A l nd ❑ A l nd ❑
Cam e ❑ Cam e ❑
Ma 9 Co ec o oca i ies y e o ius co a is. Ma 1 Co ec io oca i ies y e o ius cu ie us.
O S O E O S O E IS A S IS A S
Ke ma ecs ❑ Ke ma ecs ❑
ee Ki gs ❑ ee Ki gs ❑
C a ams ❑ C a ams ❑
S a es ❑ S a es ❑
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A i o es ❑ A i o es ❑
A l nd ❑ A l nd ❑
Cam e ❑ Cam e ❑
Ma 11 Co ec o oca i ies y e o ius ey esi. Ma 1 Co ec io oca i ies y e o ius a a .
-59- O S O E O S O E IS A S IS A S
Ke ma ecs ❑ Ke ma ecs ❑
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C a ams ❑ C a ams ❑
S a es ❑ S a es ❑
ou y ❑ ou y ❑
A i o es ❑ A i o es ❑
A l nd ❑ A l nd ❑
Cam e ❑ Cam e ❑
Ma 13 Co ec io oca i es y e o ius g acia is. Ma 1 Co ec io oca i ies y e o ius u so i.
O S O E O S O E IS A S IS A S
Ke ma ecs ❑ Ke ma ecs ❑
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C a ams ❑ C a ams ❑
S a es ❑ S a es ❑
ou y ❑ ou y ❑
A i o es ❑ A i o es ❑
A l nd ❑ A l nd ❑
Cam e ❑ Cam e ❑
Ma 15 Co ec io oca i ies y e o ius u o i. Ma 1 Co ec io oca i ies y e o ius mo a us.
- - O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ❑ 4 ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams + C a ams ❑ S a es • S a es ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ A l nd ❑ A l nd ❑ Cam e ❑ Cam e ❑
Ma 17 Co ec io oca ies y e o ius esi io es. Ma 1 Co ec io oca i ies y e o ius a icki.
O S O E O S O E IS A S IS A S Ke ma ecs ❑ Ke ma ecs ❑ ee Ki gs ❑ ee Ki gs ❑ C a ams ❑ C a ams ❑ S a es ❑ S a es ❑ ou y ❑ ou y ❑ A i o es ❑ A i o es ❑ A l nd ❑ A l nd ❑ Cam e ❑ Cam e ❑
Ma 19 Co ec io oca i ies y e o ius s e e ii. Ma Co ec io oca i ies y e o ius ow se i.
—6 — A O OMIC I E h nd x v r th n n l t x a a , y e o ius 1 1 15 17 esi io es, y e o ius 1 15 17 nt n d n th t xt r rdl f 19 5 7 3 4 , 44, 4 , , 1 19 3 3 3 53 6 m th r rr nt t t n t x n m n b r n b ldt p d n t Oc a ius d r pt n nd n t l t p ll tr t n GEOME I AE 17 o yc i us, e i o us 1 A ff x d l tt r nd t nd ge ma us, i a us 9 O o i us 1 37 p g acia is. y e o ius 1 15 19 ac y o ius 9 17 37 3 4 , , 60m A I e e a es agio oeus 37 G ACI A II AE 17 a a y es 1 aci y ae, y e o ius a am us -13 15- 37 a ameco s 37 Ac ees 11 13 17 37 ami o i, A ago us 1 a icki, y e o ius 13 15 17 31 Ac oce o s 17 e i o us 11 17 37 3 44, 4 , , 4, 6, 6 m ae io s, O o i us 1 e i us 11 37 e o a us, ysce us 1 a o i ea a, e es 1 esyc o ius 11 13 37 iceae, isso es 13 A ago us 9 1 e e e is isso es 13 17 37 A iso y c us 11 37 oma i o us 11 i os o i, a am us 1 -1 A e y o ius 11 37 u so i, y e o ius 1 1 13 15 15 19 1 37 39 48, 4 , A i icus 37 19 3 40, 4 m A E I AE 9 1 4 , 4 , , 60m i os o i, Ka oco e s 1 aus a is, y e o ius 1 15 17 19 u o i, y e o ius -1 15 1 ace i us 1 3 3 8m 1 19 3 9 3 40, 46, 2, e es 11 13 17 37 60m o o y o ius 37 a a ae, y e o ius 13 15 17 E O O I AE 17 31 3 44, 46, 0, 4, , Y O II AE 9 l b n 9 e o o us 17 8m y o i e us 17 37 AC Y E I AE 11 1 ecas 37 y o ius 9 13 17 37 y es 1 37 i e i us 1 Isc ioma us 37 Ca e ius 17 37 Sa go 1 ca i a us, y e o ius 1 13 15 Ka oco e s s e e ii, y e o ius 1 13 15 17 17 19 3 0, , , 8m 31 3 4 , 44, 4 , , Ce o 37 ac a ius, e i us 11 4, 6, 6 m C O I AE 11 1 i a us 17 37 s i i e is, a am us 1 -13 c a kei, y e o ius 1 13 15 17 15- 1 3 , 4 , m 19 5 3 40, 42, 4 , 4 , 0, E I O E A 1 S e a o y c us 17 8m i ea us, E em us 1 s i oca ae, a am us 1 - C eo i us 11 i ea us, i us 1 13 15 1 1 - 3 , co a is, y e o ius 1 13 15 1 i eige a, As yage 1 46 4 , m 19 3 3 46, 0, m i a i es 9 Sy o us 1 37 cu ie us, y e o ius 1 13 15 p r n 9 3 4 , 44, 0, 4, , m i a us 9 11 13 1 ow se i, y e o ius 13 15 15 cu ie i, i e i us 11 y e o ius -13 15-1 3 17 3 3 4 , 44, , 4, 37 6, 6 m i e sus, y es 1 u e cu a us, a am us 1 1 ysce us 11 17 37 Ma s a ius 37 15 17- 3 , 4 , m mege e i, i us 11 u e cu a us, y e o ius 9 E EM I AE 1 Me a y o ius 37 u e osus, Ca e ius 1 E is o us 1 37 M l t d 9 Eu ocimi us 37 MO Y I AE u guicu a us, e es 1 EUG OMI AE 17 mo a us, y e o ius 1 1 13 Eu ycus 37 15 17 3 3 4 , 4 , 46, , y i acma 17 ey esi, y e o ius 1 13 15 17 60m 19 7 3 4 , , 4, , m
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- 5- I ES I I / U A AI A A AA 1 e e a ia (In t h n pt r ° au e ce Α. Mou & A e e K Wa ke IS - 77- 7-9 3 19 • 1 pp $ 9 95 Oso ii ae (In t C l pt r St ph l n d • . au i e McCo IS - 77- -7 3 19 9 $1 3 A i i ae (In t C l pt r • .A. o oway IS - 77- 7 3- 3 19 • 7 pp $ 1 E io yoi ea e ce E io yi ae (Ar hn d A r • .C.M. Ma so IS - 77- 7 5- • 1 v 19 1 pp $ 9 95 5 E io yi ae (Ar hn d A r Er ph d • .C.M. Ma so IS - 77- 7 - 1 v 19 1 pp $ 9 95 y ae i ae (In t C l pt r • .G. O is IS - 77- 7 7- 1 v 19 pp $1 7 C y os igma a (Ar hn d A r - n r v • M. u o IS - 77- 7 - • 19 5 11 pp $ 9 95 Ca i o i ae (In t pt r • ames . ea IS - 77- 7 - b 19 pp $1 9 o u a (In t • S. . u e IS - 77- 7 5- b 19 5 pp $1 1 u u i e a (In t h n pt r • au e ce Α. Mou & A e e K. Wa ke IS - 77- 7 - S p 19 1 pp $3 5 11 seu ococci ae (In t pt r • .M. Co IS - 77- 791-3 7 Apr 19 7 3 pp $ 9 95 1 om i i ae (In t n pt r • A.C. a is IS - 77- 5 1-3 13 v 19 7 1 pp $39 95 13 E cy i ae (In t n pt r • .S. oyes IS - 77- 517- • 9 M 19 19 pp $ 95 1 e i o e a - nn t t d t l nd t f l - r p t x . S. ug a e • IS - 77- 51 - 3 S p 19 pp $ 9 95 15 Am osi i ae (In t n pt r pr d • I. . auma IS - 77- 535- 3 19 • 1 pp $39 95 1 e icu i ae (In t p d pt r • a s o e & C is o e Wi ki so IS - 77- 53 - Apr 19 9 9 pp $ 95 17 Myma i ae (In t n pt r • .S. oyes & Ε.W. a e i e IS - 77- 5 - Apr 19 9 1 pp $ 95 1 C a ci oi ea (In t n pt r - ntr d t n nd r v f ll r f l .S. oyes & Ε.W. a e i e • IS - 77- 5 5-5 A 19 9 9 pp $ 95 19 Ma o ea (In t th r v f p t f f n t n l rph l nd b l • G. W amsay• IS - 77- 5 1-1 • 13 n 199 • 9 pp $ 95 i io i ae (In t pt r • oy A. a iso . IS - 77- 595-1 • 13 v 199 pp $1 95
- - 1 Ma ga o i ae (In t pt r • C. . Mo a es IS - 77- 7-9 • 7 M 1991 • 1 pp $3 95 o o emou i ae (In t l pt r • I. . Mc e a IS - 77- 51 - 7 M 1991 • pp $ 95 3 Scia o i ae Me e e i ae (In t pt r th n r r v f th l h p d d • . . icke • IS - 77- 7-3 13 n 199 7 pp $ 7 95 e e i ae (In t pt r • . y e o g IS - 77- 3 - • M r 199 1 pp $3 95 5 Ce co i ae (In t pt r • K.G.A. ami o & C. . Mo a es IS - 77- 3 - 5 M 199 pp $17 95 e e io i ae (In t C l pt r t l f t p nd t t x .C. Wa • IS - 77- 39-7 13 l 199 7 pp $ 7 95 7 A a c o e i ae (In t l pt r • I. . Mc e a IS - 77- 1 - 1 b 1993 7 pp $ 7 95 a ae o Cu cu io oi ea (In t C l pt r t t v rv e a M. May. IS - 7 - 5 5- 1 n 1993 pp $55 9 C y o y c i ae (In t C l pt r C r l n d C. .C. ya • IS - 7 - 51 - 1993 3 pp $ 5 3 e ia i ae (In t p d pt r • .S. ug a e IS - 7 - 5 -7 1 M r 199 1 pp $ 5 31 a i i ae (Cr t A ph p d • K.W. u ca IS - 7 - 533- • 7 O t 199 1 pp $3 3 S eci ae (In t n pt r • A.C. a is IS - 7 - 53 - 7 O t 199 11 pp $33 5 33 Mo a i i i (In t n pt r • .A. e y IS - 7 - 53 -7 M 1995 pp $ 9 95 3 A ici ae (In t C l pt r • .G. We e & .S. C a e IS - 7 - 5 7- 1 n 1995 pp $ 5 35 Cy i ae Aca osoma i ae a e a omi ae (In t t r pt r t t r ph l d tr b t n nd b l • M. C. a i iè e IS - 7 - 93 1- 3 v 1995 11 pp $ 5 3 e o e ii ae (In t Eph r pt r • . . ow s & W. . e e s IS - 7 - 93 3-9 • 19 A 199 • 1 pp $39 5 37 C l pt r f l - r p r v nd t d nt f t n • . K imas ewski & .C. Wa • IS - 7 - 931 - 13 A 1997 199 pp $ 9 5 3 a u a ise e es ia S y omma o o a (M ll G tr p d G.M. a ke • IS - 7 - 93 -5 • n 1999 53 pp $5 39 Mo y i i (In t C l pt r C r l n d M l t n • .C. C aw IS - 7 - 93 5- • n 1999 pp $ 9 5
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h r f r f r d n l p bl t n K h tūr tēn rār n p p h h b n t bl h d t n r th wi h h h n t h n h āt r n xp rt n l d t p bl h n t - h āt n mea pā n n r r h pr h n v nt f l nt n th r n āh t h n tēn rār n l nd f n h r pr f n l n t p p n r t h h ār ai t n pt n nd pr nt t n t v r ff rt t h t h ōh ai t r n t h d t pr v d r r f r d nt f t n nd n ār r n ār r ā h ārī t āt - nf r t n th t r bl t th n n- r n pā n rāt p l t K ēn p p au a o ew ea a kāo e au a o . . d l th n n- r n nv rt- e pā a a n r r h n rā n br t nl n th v rt br t r ll n āt t rān E t n ōh t a a n d nt d nd r n f r r v r d b r r h K o o i n p p ī a a th r Ma i e au a o . . Ma i e au a o ew ea a n t h t h pā a a n r r h n n n Co i u io s r nv t d fr n p r n th āt t h th r t p l t ll nd r r M t r l fr th Arthr p d C ll t n v l bl f r t d u i u i ga K e t n n t h n t mai ā t p p E āt a a e C ntr b t r h ld d th r nt nt n th h h n r r h a a e b r f th Inv rt br t S t t l nd Arthr p d C ll t n h āt t Adv r Gr p r th th S r Ed t r b f r ā n n r ; ll n r d n ll b v n M hā ī ō t h ār t t āh t h t h au a t n t Et t Su sc i e s s ou ddr n r t au a rān mua t t t n t h t h o . , M n Wh n r nd r r h O x n ln 15 l nd ga kai- oko uka uka M t h t au a o . ., M n Wh n r nd r - S b r pt n t r A – t nd n rd r ; r h; O x n ln 15 - n nv ll b nt th h n l nd n ft r p bl t n p bl ; – pr - t n l fl r th rd r f r ll b nt fr E r n tū -h A - K -h t t t tū ; t p p p p t t r t n t tān t p p t l pr ( tl n pr nt p — t n pān n tōn ā n l d p n nd rf p t S b- tōn ā r b r n l nd nd A tr l p th nd t d nt n $ ; GS n l d d n th t (t r h t hār n K t ōp - pr Oth r b r b r p th l t d pr n t n t p n n r t t t M t $US r t v l nt t n h n r n Āh t r r n tār r n K ēt h t t t f ll n b r r v l bl nd h r t n n tār M r n n b r b r h n t bt n f ll t r l t n r t d nt ll r E t n n p p mua M h nd b r pt n nt r ff r d tr d h h n t t n p p d nt f t n p r nt t n mai h h t t E t p hēn t t h h t t n t h p p Fauna of New Zealand Ko t Aitanga Pepeke o Aotearoa
Number 39
Molytini (Insecta :Coleoptera Curculionidae Molytinae)
Robin C. Craw
O U A SUMMA Y
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C ECK IS O A A
E E E CES
KEYS O A A
A O OMIC I E h f l f th pr nt d p bl t n nd f ll r h bl It ppl d f r nd v d l nl nd n t t b p t d n b t (l n h ld b d t th p fr h h t d nl d d
p rt f th r v r d b p r ht b r pr d d r p d n n f r r b n n ( r ph l tr n r h n l n l d n ph t p n r rd n t p n nf r t n r tr v l t r th r th t th r tt n p r n f th p bl h r
Fauna of New Zealand b t p 1 fnz nd r r h nz
Cr C 1999 M l t n (In t C l pt r C r l n d M l t n Fauna of Νew ea a 39 pp
t f p bl t n br r 1999
Fauna of New Zealand, ISS 111-53 3; 39 IS - 7 - 93 5-
l nd M l t n S nn d pr v d d b St ph n n (www.bugz.org.nz). OC d t xt rr t d f r th r h bl b r v r Cr b FNZ r d t r l 1 U r xtr t t xt fr th f r th r n b t t h t n t th r n l d nt f r t xt n r nd f r tt n