Larochelle, A.; Larivière, M.-C. 2005: Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand 53, 160 pp. INVERTEBRATE SYSTEMATICS ADVISORY GROUP

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SERIES EDITOR Dr T. K. Crosby Landcare Research Private Bag 92170, Auckland, New Zealand Fauna of New Zealand Ko te Aitanga Pepeke o Aotearoa

Number / Nama 53

Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae)

A. Larochelle and M.-C. Larivière Landcare Research, Private Bag 92170, Auckland, New Zealand [email protected] [email protected]

Manaaki W h e n u a PRESS

Lincoln, Canterbury, New Zealand 2005 Copyright © Landcare Research New Zealand Ltd 2005

No part of this work covered by copyright may be reproduced or copied in any form or by any means (graphic, electronic, or mechanical, including photocopying, recording, taping information retrieval systems, or otherwise) without the written permission of the publisher.

Cataloguing in publication LAROCHELLE, ANDRÉ 1940 Apr. 10 Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae) / A. Larochelle & M.-C. Larivière, – Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 2005. (Fauna of New Zealand, ISSN 0111–5383 ; no. 53). ISBN 0-478-09369-1 I. Larivière, Marie-Claude – II. Title III. Series UDC 595.762.12(931)

Suggested citation: Larochelle, A.; Larivière, M.-C . 2005. Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand 53, 160 pp.

Prepared for publication by the series editor and the authors using computer-based text processing, layout, and printing at Landcare Research, Private Bag 92170, Auckland, New Zealand. Frontispiece digital image produced by M.-C. Larivière using the synchroscopy system Auto-Montage®.

M~ori text by H. Jacob, Auckland.

Published by Manaaki Whenua Press, Landcare Research, P.O. Box 40, Lincoln, Canterbury, N.Z. Website: http://www.mwpress.co.nz/

Printed by PrintLink Ltd, Wellington

Front cover: Tuiharpalus moorei new species (Illustrator: D. W. Helmore).

Publication of the Fauna of New Zealand series is the result of a research investment by the Foundation for Research, Science and Technology under contract number C09X0202. Fauna of New Zealand 53 5

POPULAR SUMMARY HE WHAKARAPOPOTOTANGA

Class Insecta Order Coleoptera Family Carabidae Subfamily Harpalinae Tribe Harpalini

Harpaline ground beetles The tribe Harpalini belongs to the subfamily Harpalinae Illustration / Whakaahua: Lecanomerus sharpi (Csiki) (Coleoptera: Carabidae), which contains over 19 000 taxa (Illustrator / Kaiwhakaahua: D. W. Helmore). worldwide. Molecular sequence data indicate Harpalinae evolved in the Cretaceous Period (140–65 million years Ng~ p§tara Harpaline noho papa ago). The Harpalini form a diverse group, including over 240 NÇ te wh~nau wh~iti Harpalinae (Coleoptera: Carabidae) genera and subgenera, and approximately 2000 species a ng~i Harpalini. Puta noa i te ao, 1900 Çna rÇpã. E ai ki distributed in all biogeographic regions of the world. The ng~ raraunga raupapa r~poi ngota, nÇ te Takiw~ Creta- present faunal review records 20 genera and 57 species for ceous tÇna kunenga mai (i te 140–65 miriona tau ki muri). New Zealand. This should constitute nearly all the fauna. He iwi matahuhua tonu a ng~i Harpalini – he nui atu i Compared with larger or warmer regions of the world, te 240 ng~ puninga me ng~ puninga iti, he ~hua 2000 ng~ e.g., Australia, which has a largely undescribed fauna with momo, kei ng~ takiw~ papawhenua-koiora katoa o te ao. I over 160 known species, the New Zealand fauna may t‘nei tirohanga hou, kua tuhia he mauranga mÇ ng~ puninga appear relatively small, but New Zealand is a very special e 20 me ng~ momo e 57 i Aotearoa. Ko te whakaaro ia, he place – a biodiversity ‘hot-spot’ – with 75% of species (42 ruarua noa iho ng~ momo k~ore i mau i t‘nei tatauranga. out of 57 species) and 55% of genera (11 out of 20 genera) Ina whakaritea ki ng~ whenua rahi ake, mahana ake o found nowhere else in the world. The remaining fauna that te ao (hei tauira, ar~ a Ahitereiria me Çna momo 160 e are not endemic to this country are made up of overseas mÇhiotia ana, tae atu ki te maha noa atu k~ore anÇ i species introduced mainly from Australia. No native species whakaahuatia ~-kupu) t‘r~ ka whakaarohia he torutoru is shared with Australia, although three native genera occur noa iho ng~ momo o Aotearoa. Engari he whenua ahurei on both sides of the Tasman Sea, which suggests the New tonu a Aotearoa, in~ r~, ko t‘tahi 75% o ng~ momo (e 42 o Zealand harpaline lineages have evolved mostly in isolation ng~ momo e 57) me t‘tahi 55% o ng~ puninga (11 o ng~ following the breakup of eastern Gondwanaland. puninga e 20) i Aotearoa nei, k~ore e kitea i whenua k‘. Ko Harpalini are rather stout-bodied ground beetles with ‘r~ atu momo ehara nÇ konei taketake ake, he r~waho, ko relatively short mandibles and other appendages, and a te nuinga i manene mai i Ahitereiria. Karekau he momo body length of 3–20 mm. They are usually darkly coloured, m~ori o Aotearoa kei Ahitereiria anÇ e noho ana, engari e have only one hair-bearing puncture above each eye, no toru ng~ puninga m~ori kei ng~ whenua e rua nei. E tohu such puncture at the posterior angles of the pronotum, and ana t‘nei t‘r~ tonu pea i kukune motuhake mai ng~ t~tai elytra that are rounded, not twisted, at the sides near the harpaline o Aotearoa i muri i te wehewehenga o te pito apex. Some species living in caves or exhibiting strong r~whiti o Te Uri M~roa. burrowing habits are characterised by paler bodies and He ~hua porotaka ng~ tinana o ng~ p§tara noho papa reduced eyes. Harpalini, he ~hua poto ng~ kauwae me ‘r~ atu w~hanga As observed in many other carabids, harpaline ground toro whakawaho o te tinana. Ko te roa o te tinana, mai i te beetles are taxonomically diverse, generally abundant in 3 ki te 20 mm. He uriuri te tae o te nuinga, kotahi anake te the field, and demonstrate ecological preferences and a m~rua whai weu i runga ake o ia karu, k~ore he m~rua flexible set of responses to environmental factors. Because p‘nei i ng~ koki whakamuri o te papatua pohomua. He of these features, the relative ease with which their ~hua kÇpuku ng~ pãkoro parirau i ng~ kaokao, i te takiw~ (continued overleaf) (haere tonu) 6 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) populations may be sampled by reliable quantitative o te pito, k~ore e kÇrino. Heoi, ar~ ‘tahi momo noho ana, methods (e.g., pitfall-trapping), and their potential use as kari rua r~nei k~ore e tino uriuri ng~ tinana, he iti ake anÇ bioindicators and biocontrol agents, they represent an hoki ng~ karu. attractive study group for biologists investigating P‘r~ anÇ i te maha tonu o ng~ carabid, he matahuhua evolutionary and ecological hypotheses. ng~ whakarÇpãtanga o ng~ p§tara noho papa harpaline, he As a result, Harpalini are well represented in New huhua anÇ t~ r~tou noho ki te taiao. He rerek‘ ng~ k~inga Zealand entomological museums and collections – over noho e pai ana ki t‘n~, ki t‘n~, he t~wariwari anÇ te ~hua 5000 specimens were studied for this project. But despite o t~ r~tou aro atu ki ng~ ~huatanga taiao. N~ ‘nei such high interest, no taxonomic revision of this group has ~huatanga, n~ te m~m~ anÇ ki te t§pako i ng~ taupori i been produced until now. runga i ng~ tikanga ine ~-rahi tÇtika (p‘nei i te hopu ki te Before the present revision, 13 genera and 36 species tomo), me tÇ r~tou pai anÇ pea hei tohu koiora, hei kaipatu of Harpalini were known from New Zealand, but the koiora r~nei, he rÇpã t‘nei e arohia nuitia ana e ng~ tohunga authors have found a number of species had been described koiora e whakam~tau ana i ng~ whakapae mÇ te kunenga more than once under different names, and 23 species and me te taupuhi kaiao. 5 genera are new to science. Me te aha, he autaia tonu te maha o ng~ Harpalini e The geographical distribution of native species was puritia ana ki ng~ whare pepeke me ng~ kohinga pepeke i undocumented before this study. We now have a better Aotearoa – he nui ake i te 5000 ng~ tauira i ~ta tirohia mÇ knowledge of their distribution patterns. The authors have t‘nei rangahautanga. Engari ahakoa te aro nui ki t‘nei iwi, found, for example, that several species are restricted to k~tahi anÇ ka tirohia anÇ te whakarÇpãtanga o ng~ hanga specific areas of New Zealand – the South Island northwest nei. and the far north of the New Zealand appearing to have I mua i t‘nei o ng~ tirohanga, 13 ng~ puninga, e 36 ng~ been reservoirs, in geological time, of much of the genetic momo i mÇhiotia i Aotearoa nei. Engari ko t~ ng~ kaituhi i diversity in New Zealand Harpalini, with several species kite ai, ar~ ‘tahi momo i tapaina ki ng~ ingoa e rua, neke currently restricted to these regions. Of the two main atu r~nei, me Çna anÇ kupu whakaahua i te taha. }, e 23 islands of New Zealand, the North Island has the greatest ng~ momo, e 5 ng~ puninga k~ore i mÇhiotia i te ao pãtaiao number of species (35 compared with 31 for the South i mua atu i t‘nei. Island), and only 4 native species (Allocinopus sculpticollis, Waihoki, k~ore i tuhia te tohanga o ng~ momo m~ori i Triplosarus novaezelandiae, Syllectus anomalus, mua atu i t‘nei rangahautanga. N~ t ‘nei mahi rangahau kua Euthenarus puncticollis) are shared between the two islands. kaha ake te mÇhio ki Ç r~tou tauira tohatoha. Hei tauira, i Three genera (Gaioxenus, Parabaris, and Kupeharpalus) kitea ar~ ‘tahi momo maha tonu e noho motuhake ana ki are found only on the North Island, whereas two genera ‘tahi rohe wh~iti o Aotearoa – ko te uru-m~-raki o Te (Hakaharpalus and the cave-dwelling Pholeodytes) are Waipounamu me Muriwhenua ‘tahi tino m~t~puna, ~-w~ confined to the northwest of the South Island. Two genera papawhenua nei, o te matahuhuatanga o te huinga ira o are restricted to the Three Kings Islands (Maoriharpalus, ng~ Harpalini o Aotearoa. He maha hoki ng~ momo k~ore Kiwiharpalus). There is no genus endemic to the Chatham e kitea i waho atu o ‘nei takiw~. He maha ake ng~ momo i Islands. Stewart Island also has no endemic taxa, but shares Te Ika a M~ui (e 35 ng~ momo), t‘n~ i Te Waipounamu (e 2 native species: Triplosarus novaezelandiae (with North 31 ng~ momo), ~, e 4 anake ng~ momo m~ori (Allocinopus Island and South Island), Euthenarus brevicollis (with South sculpticollis, Triplosarus novaezelandiae, Syllectus Island). anomalus, Euthenarus puncticollis) kei ng~ moutere e rua Over 50% of native species (25 out of 42 species) are nei. E 3 ng~ puninga (Gaioxenus, Parabaris me known from 10 populations or fewer. All but one of these Kupeharpalus) kei Te Ika a M~ui anake, e 2 very special species are new to science, and all are of (Hakaharpalus me Pholeodytes – he momo noho ana) kei potential conservation concern. They are usually te uru-m~-raki o Te Waipounamu anake. E 2 ng~ puninga taxonomically highly distinctive species with low dispersal k~ore e kitea i waho atu o Manawa-t~whi (Maoriharpalus, power, often geographically localised in threatened habitats, Kiwiharpalus). Karekau he puninga e kitea ana i R‘kohu and represented in collections by relatively few specimens anake. Waihoki, k~ore he rÇpã e kitea ana i Rakiura anake, collected over many decades, which may indicate rare or engari e 2 Çna momo m~ori, ko: Triplosarus novaezelandiae highly specialised species. (kei R‘kohu, kei Te Ika a M~ui me Te Waipounamu), me No formal detailed study of the natural history of Euthenarus brevicollis (kei R‘kohu me Te Waipounamu). individual species of New Zealand Harpalini has ever been He nui ake i te 50% o ng~ momo m~ori (e 25 o ng~ conducted, although Larochelle and Larivière (2001, Fauna momo e 42), nÇ roto i ng~ taupori 10, iti ake r~nei. E 24 o of New Zealand 43) summarised information available from ‘nei momo, k~tahi anÇ ka mÇhiotia i te ao pãtaiao, ~, ko te the literature, material in entomological collections, personal katoa me ~ta tiaki ka tika kei koreh~h~ haere. Ko te nuinga, communications from carabid collectors, and their own he m~m~ ki te wehewehe, t‘tahi i t‘tahi, i te nui o ng~ personal field observations. rerek‘tanga, k~ore e kaha te marara haere, e noho wh~iti Most native species are flightless, having vestigial ana ki ng~ ripoinga mÇrearea, ~, he ruarua ng~ tauira o t‘n~, membranous wings (reduced to small wing buds), and live o t‘n~ kua kohia i roto i ng~ tekau tau, e tohu ana he momo within the confines of native habitats, mostly forests onge tonu pea, he tino wh~iti r~nei Ç r~tou k~inga noho. (continued overleaf) (haere tonu) Fauna of New Zealand 53 7

(especially along streams) and wet habitats, also tussock K~ore anÇ i ~ta rangahaua ng~ h§tori m~ori o t‘n~, o grasslands and caves. Most Harpalini species are moisture t‘n~ momo o ng~i Harpalini i Aotearoa, engari i loving and live at the surface of the soil and in leaf litter; whakar~popotohia e Larochelle r~ua ko Larivière (2001, they also live in caves, and occasionally on plants and Fauna of New Zealand 43) ng~ p~rongo kua tuhia, ng~ trees. Dispersal in most native species is achieved by rauemi i ng~ kohinga pepeke, ng~ whakawhitinga kÇrero running over the ground; most species are moderate runners, ki ng~ kaikohikohi carabid, me Ç r~ua ake kitenga i te taiao. except for the long-legged, fast-running cave species He rere kore te nuinga o ng~ momo m~ori, he tumu (Syllectus, Pholeodytes). In general, Harpalini have relatively parirau noa iho, he mea hanga ki te kiriuhi, e toe mai ana. short legs and, sometimes, strongly reduced eyes, indicative Ka mutu, noho ai te nuinga ki ng~ ripoinga m~ori, ar~, ki of strong burrowing habits. ng~ ngahere m~ori (ko ng~ tahataha o ng~ kÇawa t‘tahi tino All adventive species are able to fly, having long or k~inga) tae atu ki ng~ k~inga kÇreporepo, ng~ whenua fully developed membranous wings, and live mostly in p~t§t§ taranui me ng~ ana. He pai ki te nuinga t‘nei mea te highly modified environments (often around human haukã, ~, t‘r~ ka noho ki te mata tonu o te whenua, ki ng~ dwellings), except for Haplanister crypticus, which has rau popo. Ar~ anÇ ‘tahi ko te ana tÇ r~tou w~hi noho, ~, he also managed to invade native forests. torutoru ng~ momo ka piri tahi ki ng~ r~kau me ‘r~ atu The collecting season of newly emerged adults suggests tipu. Ko te tikanga t§tari a te nuinga o ng~ momo m~ori, ko Harpalini species may mate in the spring or autumn. For te takahi i te nuku o te whenua; he ~hua tere te haere a te most species, adults are active during all months of the nuinga, engari he tere tonu te momo noho ana, waewae roa year, but are generally less active during cooler months. (Syllectus, Pholeodytes). He poto ng~ waewae o te nuinga No data are available on the feeding preferences of o ng~i Harpalini, ~, he tino ngoikore ng~ karu, e tohu ana Harpalini native to New Zealand. Larochelle (1990, Food he kari rua, he noho rua t~ r~tou mahi. of carabid beetles of the World) showed that on a world Katoa ng~ momo r~waho, he mÇhio ki te rere, ~, he basis Harpalini feed on both animal and vegetable matter, roa, he pakari r~nei Ç r~tou parirau kiriuhi. Noho ai te but tend to favour the latter. The mandibles of nuinga ki ng~ taiao kua kaha rawekehia e te ringa tangata Hakaharpalus, Kiwiharpalus, Syllectus, Pholeodytes, and (he maha e noho tata ana ki ng~ whare), h~unga anÇ te Maoriharpalus are unusually long in native Harpalini, which Haplanister crypticus. Kua urutomo anÇ t‘nei n~ i ng~ may suggest a specialised type of feeding. In addition, the ngahere m~ori. strongly notched labrum (upper mouth part at base of Ko te w~ e kohikohia ai ng~ p§tara pakeke k~tahi anÇ mandibles) of Maoriharpalus is reminiscent of, although ka puta ake ki te ao e tohu ana t‘r~ pea ko te kÇanga, ko te not necessarily equivalent to, the condition observed in the ngahuru r~nei te w~ e whakaputa uri ai a ng~i Harpalini. tribe Licinini, where species feed on hard-bodied MÇ te nuinga o ng~ momo, e oreore ana ng~ mea pakeke i invertebrates, e.g., snails. ng~ marama katoa o te tau, engari ka ~hua ngoikore ake i This faunal review was written with a wide audience in ng~ marama makariri ake. mind. It aims to provide an inventory of New Zealand Karekau he raraunga mÇ ng~ kai e pai ana ki ng~ taxa, a concise treatment of their taxonomy, easy-to-follow Harpalini m~ori. Heoi, n~ Larochelle (1990, Food of carabid identification keys, and several illustrations and maps, as beetles of the World) i whakaatu kai ai ng~ Harpalini o te well as a summary of all available information on species ao i te kiko me te ota, engari ko te ota pea t~ r~tou tino kai. distribution, ecology, biology, and dispersal power. It is He roa ake ng~ kauwae o Hakaharpalus, Kiwiharpalus, one step in the authors’ goal to to reach an overall Syllectus, Pholeodytes, me Maoriharpalus i ‘r~ o te nuinga understanding of the carabid fauna within a reasonable time o ng~ momo m~ori, e tohu ana pea he rerek‘ ng~ momo kai frame, and to make relatively large amounts of information kainga ai e r~tou. I tua atu i t‘r~, ar~ ‘tahi k~niwha nui i te available for practical use by a wide range of end-users. It ngutu o runga o Maoriharpalus, e ~hua rite ana ki t‘r~ e is hoped this kind of faunal taxonomy will provide both a kitea ana i te iwi Licinini. Ko t~ t‘r~ iwi, he kai i ng~ hanga solid foundation for studies of other types and the baseline tuar~-kore, tinana m~rÇ, p‘r~ i te ngata. information required by systematists, identifiers, ecologists, He mea tuhi t‘nei tirohanga hou kia m~rama ai ng~ and other biologists, as well as by biosecurity and kÇrero ki te tokomaha. E whai ana kia takoto mai he r~rangi conservation managers. o ng~ rÇpã e noho ana ki Aotearoa nei, he whakam~rama poto o Ç r~tou t~tai hono, he ara tautohu m~m~, he whakaahua, he mahere, tae atu ki t‘tahi whakar~- Contributor André Larochelle was born and educated in popototanga o ng~ p~rongo e w~tea ana mÇ te tohanga o Québec, graduating in 1974 with a Brevet d’Enseignement ~ spécialisé from the Université du Québec à Montréal. He ng momo, te taupuhi kaiao, te koiora, me te kaha ki te § ~ Ç ‘ taught ecology at the Collège Bourget, Rigaud, Québec, up t tari haere i a r tou an . He takahanga noa t nei i roto i te ~ ~ ~ to 1990. With the encouragement of the late carabid spe- wh inga roa a ng kaituhi kia wh nui noa ake te cialist Carl H. Lindroth, André very quickly becames inter- m~ramatanga ki ng~ carabid i roto i te tekau tau pea e tã ested to the study of ground beetles. From 1975 to 1979 he mai nei, kia hora he p~rongo huhua hei ~whina i ng~ t~ngata was the co-editor of two entomological journals, Cordulia tokomaha i roto i ~ r~tou kaupapa maha. Ko te tãmanako and Bulletin d’inventaire des insectes du Québec. From ia, kia noho t‘nei tãmomo whakarÇpãtanga hei tã~papa (continued overleaf) (haere tonu) 8 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

1986 to 1992, he was honorary curator to the Lyman Ento- mÇ ng~ mahi rangahau i ‘tahi atu pepeke, hei puna kÇrero mological Museum and Research Laboratory, McGill Uni- hoki m~ ng~ kait~tai whakapapa, ng~ kaitautohu, ng~ versity, Québec. In 1992, André moved to New Zealand to tohunga taupuhi kaiao, me ‘tahi atu ringa koiora, tae atu ki work as a research scientist. Currently, he is a Research ng~ kaiwhakahaere haumaru koiora, tiaki taiao. Associate with the New Zealand Arthropod Collection, Landcare Research, Auckland. André has written over 400 papers on the distribution, ecology, biology, and dispersal power of North American carabids and other insects (in- cluding two handbooks on the Heteroptera of Québec). In I wh~nau mai t‘r~ atu o ng~ kaituhi, a André Larochelle, 1993 he was co-author of a “Catalogue of Carabidae of i Québec. I reira anÇ ia e kura ana, ~, nÇ te tau 1974 ka whakawhiwhia ki tana tohu Brevet d’Enseignement spécialisé, mai i te Whare W~nanga o Québec ki Montreal. Taka mai ki te tau 1990, e whakaako ana ia i te m~tauranga taupuhi kaiao i te K~reti Bourget, i Rigaud, Québec. N~ ng~ akiaki a t‘r~ tohunga carabid kua riro nei i te tirohanga kanohi, a Carl H. Lindroth, ka tere tupu tana hiahia ki te rangahau i ng~ p§tara noho papa. Mai i te 1975 ki te 1979 ko ia t‘tahi o ng~ ‘tita o ‘tahi hautaka m~tai pepeke e rua, ar~, o Cordulia me te Bulletin d’inventaire des insectes du Québec. Mai i te 1986 ki te 1992, ko ia te kaitiaki utu-kore o te Whare Pupuri, Rangahau Pepeke o Lyman, i te Whare W~nanga o McGill, i Québec. I te tau 1992, ka neke mai a André ki Aotearoa, ka mahi hei kaipãtaiao rangahau. I t‘nei w~, e noho ana ia hei Kairangahau i te Kohinga Angawaho o Aotearoa, i Manaaki Whenua ki T~maki- makau-rau. He nui ake i te 400 ng~ kÇrero kua tuhia e André mÇ te tohanga, te taupuhi kaiao, te koiora, me te t§taringa o ng~ carabid me ‘tahi atu aitanga pepeke o Amerika ki te Raki (tae atu ki ‘tahi pukapuka e rua e whakaahua ana i ng~ Heteroptera o Québec). I te tau 1993 ko ia t‘tahi o ng~ kaituhi i te “R~rangi o ng~ Carabidae o Amerika ki te raki o M‘hiko”. Ka whakaputaina e r~ua ko tana hoa wahine, a Marie-Claude, “Ng~ H§tori M~ori o ng~ Carabidae” mÇ taua takiw~ anÇ (2003), t‘tahi r~rangi o ng~ Carabidae (2001) me ng~ Heteroptera (2004) o Aotearoa. Ko te aronga nui o ~na rangahau i t‘nei w~, ko te America north of Mexico”. With his wife, Marie-Claude, ~hua me te whakarÇpãtanga o ng~ p§tara noho papa o he published “A Natural History of Carabidae” for the Aotearoa. Ko t‘tahi w~hanga nui o ‘nei, ko tana pukapuka same region (2003) as well as a catalogue of New Zealand ~whina i te tautohunga o ng~ iwi me ng~ puninga Carabidae Carabidae (2001) and Heteroptera (2004). His current main o Aotearoa, taihoa nei ka puta. research interest is the faunistics and taxonomy of New Zealand ground beetles, which involves a soon-to-be-pub- lished identification guide to the tribes and genera of Carabidae from New Zealand. I wh~nau mai a Marie-Claude Larivière i Québec. I reira anÇ ia e rapu ana i te m~tauranga ~, riro noa i a ia tana Contributor Marie-Claude Larivière was born and edu- Tohu T~kutatanga mai i te Whare W~nanga o McGill, i te cated in Québec, graduating with a Ph.D. in systematic tau 1990, ko ng~ whakapapa pepeke te kaupapa. MÇ te entomology from McGill University in 1990. For the fol- rua tau i muri mai, kei Agriculture Canada, i Ottawa, ia e lowing 2 years she did postdoctoral research at Agriculture wh~wh~ ana i ‘tahi atu rangahautanga. I te tau 1992, ka Canada, Ottawa. In 1992, Marie-Claude moved to New neke mai a Marie-Claude ki Aotearoa, ka mahi hei kait~tai Zealand to work as a full-time Hemiptera biosystematist i ng~ whakapapa o ng~i Hemiptera i Manaaki Whenua. with Landcare Research. From 1994 to 1997 she led the N~na i ~rahi Te T~tainga o ng~ Whakapapa o ng~ Aitanga Biosystematics of New Zealand Land Invertebrates pro- Tuar~-Kore a T~ne mai i te tau 1994 ki te 1997, me te gramme, and from 1999 to 2004, the Koiora-BioAssist™ kaupapa Koiora-BioAssist™ (Te Aromatawai i ng~ Koiora project (Biodiversity Assessment using Information Tech- i runga i te Whakamahi i te Hangarau MÇhiohio me te nology and Taxonomy). Marie-Claude is the author of over WhakarÇpãtanga) mai i te tau 1999 ki te 2004. He neke 70 papers and monographs on the taxonomy, distribution, atu i te 70 ng~ tuhinga kua puta i a ia e p~ ana ki te (continued overleaf) (haere tonu) Fauna of New Zealand 53 9 and natural history of Hemiptera and Carabidae whakarÇpãtanga, te tohanga me te h§tori m~ori o ng~ Hemi- (Coleoptera), including four Fauna of New Zealand contri- ptera me ng~ Carabidae (Coleoptera), tae atu ki ‘tahi butions (Hemiptera—Cixiidae and Pentatomoidea revisions, putanga e wh~ o Te Aitanga Pepeke o Aotearoa. He tuhinga catalogues—Carabidae and Heteroptera ). She has also anÇ kua puta i a ia mÇ ng~ Orthoptera me ng~ Carabidae o published on North American Orthoptera and Carabidae. Amerika ki te Raki. Kua mahi tahi anÇ r~ua ko tana hoa Many of her publications were written in collaboration t~ne, a André, ki te whakaputa i ng~ tuhinga huhua. Ko te with her husband André with whom she hopes to soon tãmanako, taihoa ka puta i a r~ua he tuhinga hou mÇ ng~ publish new works on New Zealand Hemiptera and Hemiptera me ng~ Carabidae o Aotearoa. Kei te ng~kaunui Carabidae. Marie-Claude has a keen interest in biodiversity anÇ a Marie-Claude ki te p~rongo-koiora, tae atu ki te informatics, especially digital taxonomy, computer imaging, whakarÇpãtanga ~-mati, te t~rai whakaahua ki te rorohiko, interactive identification, and web-publishing. te tautohu i runga i te mahi p~hekoheko, me te p~nui kÇrero ki te pae tukutuku.

Translation by H. Jacob T~maki-makau-rau / Auckland

DEDICATION “Think where man’s glory most begins and ends And say my glory was I had such friends” W. B. Yeats 1865-1939: The Municipal Gallery Re-visited (1939) We are glad to dedicate this revision to our colleague Barry P. Moore (Research Associate, Australian National Insect Collection, Canberra) in acknowledgement of his continued friendship towards New Zea- land coleopterists and his generously provided expertise on New Zealand carabids. Over the years Barry has kindly identified ground-beetles for the New Zealand Arthropod Collection and private collectors, at a time when a large proportion of the fauna remained undescribed. His publications on the carabids of New Zealand (e.g., 1980, Anillina; 1996, Haplanister crypticus) and Australia (e.g., 1987, Australian catalogue) have provided a solid foundation for our New Zealand catalogue (Larochelle & Larivière, 2001) and future taxonomic revisions. In the preparation of the Harpalini revision, Barry has generously given us much encouragement and support in the identification of adventive species as well as useful comments on some difficult taxonomic problems. 10 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Frontispiece: Triplosarus novaezelandiae (Laporte de Castelnau, 1867) (photograph prepared by M.-C. Larivière, Landcare Research) Fauna of New Zealand 53 11

ABSTRACT The New Zealand Harpalini fauna (Coleoptera: Carabidae: Harpalinae) is revised. Twenty genera and fifty-seven species are recognised. Five new genera are de- scribed: Maoriharpalus new genus and Tuiharpalus new genus (Anisodactylina), Hakaharpalus new genus and Kupeharpalus new genus (Pelmatellina), and Kiwiharpalus new genus (Stenolophina). The species Parabaris gourlayi Britton, 1964 is transferred to Tuiharpalus new genus. The original combination Harpalus australasiae Dejean, 1829 is reinstated. Tachys (?) cavelli Broun, 1893 is trans- ferred to the genus Hakaharpalus. Twenty-three new species are described: Anisodactylina — Allocinopus belli new species, A. bousqueti new species, A. wardi new species, Maoriharpalus sutherlandi new species, Parabaris hoarei new species, P. lesagei new species, Tuiharpalus clunieae new species, T. crosbyi new species, T. hallae new species, T. moorei new species; Pelmatellina — Hakaharpalus davidsoni new species, H. maddisoni new species, H. patricki new species, H. rhodeae new species, Kupeharpalus barrattae new species, K. embersoni new species, K. johnsi new species, Lecanomerus marrisi new species, Syllectus gouleti new species; Stenolophina — Kiwiharpalus townsendi new spe- cies, Pholeodytes helmorei new species, P. nunni new species, and P. palmai new species . Eight new synonymies are established (valid name listed after equal sign): Allocinopus ocularius Broun, 1908 = Allocinopus sculpticollis Broun, 1903; Allocinopus castaneus Broun, 1912 = Allocinopus smithi Broun, 1912; Hypharpax abstrusus Bates, 1878 = Hypharpax australis (Dejean, 1829); Lecanomerus fallax Broun, 1880 = Lecanomerus insignitus Broun, 1880; Lecanomerus fuliginosus Broun, 1880, Lecanomerus pallipes Broun, 1894, and Lecanomerus incertus Broun, 1914 = Lecanomerus latimanus Bates, 1874; Syllectus spelaeus Britton, 1964 = Syllectus magnus Britton, 1964. Lectotypes are designated for the following taxa: Allocinopus smithi Broun, 1912; A. castaneus Broun, 1912; A. latitarsis Broun, 1911; Euthenarus brevicollis Bates, 1874; E. puncticollis Bates, 1874; Gaioxenus pilipalpis Broun, 1910; Harpalus antarcticus Laporte de Castelnau, 1867, H. novaezelandiae Laporte de Castelnau, 1867, Hypharpax abstrusus Bates, 1878; Lecanomerus fallax Broun, 1880; L. obesulus Bates, 1878; Syllectus anomalus Bates, 1878; Triplosarus fulvescens Bates, 1874. Four adventive species are re- corded for New Zealand for the first time: Gnathaphanus melbournensis (Laporte de Castelnau, 1867); Notiobia quadricollis (Chaudoir, 1878); Euthenarus bicolor Moore, 1985; E. promptus (Erichson, 1842). A concise revision of the taxonomy of all taxa is provided. Subtribes, genera, and species are keyed. Descriptions are provided with illustrations emphasising the most important diagnostic features of the external morphology and male genitalia. Information is given on synonymy, type data, material examined, geographic distribution, ecology, biology, dispersal power, and collecting techniques. The composition of the New Zealand Harpalini fauna, with endemism levels of 55% for genera and 75% for species, and its affinities with Australia, New Caledonia, Lord Howe Island, and Norfolk Island are analysed and discussed. Over 50% of native taxa (25 out of 42 species) are known from 10 populations or fewer and may be of potential conservation concern.

Keywords: Coleoptera, Carabidae, Harpalini, new genera, new species, adventive species, taxonomy, keys, distribution, ecology, biology, dispersal power, fauna.

Larochelle, A.; Larivière, M.-C. 2005: Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand 53, 160 pp.

Received: 28 June 2004. Accepted 13 October 2004. 12 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

CHECKLIST OF TAXA Genus Notiobia Perty, 1830 A ...... 41 Subgenus Anisotarsus Chaudoir, 1837 A ...... 41 Note: Valid taxa are listed alphabetically (A=Adventive, Diaphoromerus Chaudoir, 1843 E=Endemic, N=Native, but not endemic to New Zealand). Eurytrichus LeConte, 1848 Stilbolidus Casey, 1914 Family CARABIDAE quadricollis (Chaudoir, 1878) A first New Zealand record ...... 41 Subfamily HARPALINAE Genus Parabaris Broun, 1881 E ...... 42 Tribe HARPALINI ...... 24 atratus Broun, 1881 E ...... 43 E Subtribe ANISODACTYLINA ...... 26 hoarei new species ...... 44 E Genus Allocinopus Broun, 1903 E ...... 27 lesagei new species ...... 43 E angustulus Broun, 1912 E ...... 29 Genus Triplosarus Bates, 1874 ...... 45 E belli new species E ...... 30 novaezelandiae (Laporte de Castelnau, 1867) .. 45 bousqueti new species E ...... 31 fulvescens Bates, 1874 E latitarsis Broun, 1911 E ...... 32 Genus Tuiharpalus new genus ...... 46 E sculpticollis Broun, 1903 E ...... 33 clunieae new species ...... 48 E ocularius Broun, 1908 new synonym crosbyi new species ...... 47 E smithi Broun, 1912 E ...... 28 gourlayi (Britton, 1964) new combination ...... 48 E castaneus Broun, 1912 new synonym hallae new species ...... 49 E wardi new species E ...... 31 moorei new species ...... 50 Genus Anisodactylus Dejean, 1829 A ...... 34 Subtribe HARPALINA ...... 50 A Subgenus Anisodactylus Dejean, 1829 A ...... 34 Genus Harpalus Latreille, 1802 ...... 51 A binotatus (Fabricius, 1787) A ...... 34 Subgenus Harpalus Latreille, 1802 ...... 51 Amblystus Motschulsky, 1864 Genus Gaioxenus Broun, 1910 E ...... 35 affinis (Schrank, 1781) A ...... 51 pilipalpis Broun, 1910 E ...... 35 aeneus Fabricius, 1775, Carabus homonym Genus Gnathaphanus Macleay, 1825 A ...... 36 tardus (Panzer, 1797) A ...... 52 Pachauchenius Macleay, 1864 Subgenus (Uncertain) Mirosarus Bates, 1878 australasiae Dejean, 1829 A reinstated ...... 53 melbournensis (Laporte de Castelnau, 1867) A first Subtribe PELMATELLINA ...... 54 New Zealand record ...... 36 Genus Hakaharpalus new genus E ...... 54 paroensis Laporte de Castelnau, 1867, Harpalus cavelli (Broun, 1893) E new combination ...... 57 marginicollis Laporte de Castelnau, 1867, davidsoni new species E ...... 56 Harpalus maddisoni new species E ...... 56 adelaideae Laporte de Castelnau, 1867, Harpalus patricki new species E ...... 55 planipennis Macleay, 1871, Harpalus rhodeae new species E ...... 57 angustatus Macleay, 1871, Harpalus Genus Kupeharpalus new genus E ...... 57 aeneonitens Macleay, 1871, Harpalus barrattae new species E ...... 58 gayndahensis Macleay, 1871, Harpalus embersoni new species E ...... 59 atroviridis Macleay, 1871, Harpalus johnsi new species E ...... 59 insularis Bates, 1878, Mirosarus Genus Lecanomerus Chaudoir, 1850 N ...... 60 Genus Hypharpax Macleay, 1825 N ...... 37 Thenarotes Bates,1878 Sagraemerus Redtenbacher, 1868 Odontagonum Darlington, 1956 antarcticus (Laporte de Castelnau, 1867) E ...... 38 atriceps (Macleay, 1871) A ...... 62 australis (Dejean, 1829) A ...... 39 javanus Jedli…ka, 1964, Acupalpus inornatus Germar, 1848, Harpalus insignitus Broun, 1880 E ...... 63 coxii Laporte de Castelnau, 1867, Harpalus fallax Broun, 1880 new synonym abstrusus Bates, 1878 new synonym latimanus Bates, 1874 E ...... 64 parvus Chaudoir, 1878 fuliginosus Broun, 1880 new synonym Genus Maoriharpalus new genus E ...... 40 pallipes Broun, 1894 new synonym sutherlandi new species E ...... 40 incertus Broun, 1914 new synonym Fauna of New Zealand 53 13

marrisi new species E ...... 66 Subtribe Harpalina ...... 50 obesulus Bates, 1878 E ...... 63 Subtribe Pelmatellina ...... 54 sharpi (Csiki, 1932) E ...... 65 Subtribe Stenolophina ...... 72 marginatus Sharp, 1883 homonym Bibliography ...... 83 verticalis (Erichson, 1842) A ...... 67 Appendix A: Glossary of technical terms ...... 89 insidiosus Chaudoir, 1850 Appendix B: Geographical coordinates of main localities flavocinctus Blackburn, 1888 ...... 91 occidentalis Sloane, 1898 Addendum ...... 93 vestigialis (Erichson, 1842) A ...... 67 Illustrations ...... 94 mastersii Macleay, 1871, Acupalpus Map 1. The New Zealand subregion ...... 140 stenopus Broun, 1886 Map 2. Area codes and collecting localities from mainland nitidus Blackburn, 1891 New Zealand: North Island ...... 141 labralis Broun, 1914 Map 3. Area codes and collecting localities from mainland Genus Syllectus Bates, 1878 E ...... 68 New Zealand: South Island and Stewart Island . 142 anomalus Bates, 1878 E ...... 69 Map 4. Total number of known taxa by areas ...... 143 gouleti new species E ...... 71 Map 5. Number of known New Zealand endemics by magnus Britton, 1964 E ...... 70 areas ...... 144 spelaeus Britton, 1964 new synonym Map 6. Number of native taxa known to be restricted to Subtribe STENOLOPHINA ...... 72 single areas ...... 145 Genus Egadroma Motschulsky, 1855 A ...... 72 Map 7. Number of known adventive taxa by areas . 146 picea (Guérin-Méneville, 1830) A ...... 73 Species distribution maps ...... 147 dingo Laporte de Castelnau, 1867, Harpalus Taxonomic index ...... 154 politus Macleay, 1871, Stenolophus sexualis Fauvel, 1882, Stenolophus Genus Euthenarus Bates, 1874 N ...... 73 ACKNOWLEDGMENTS bicolor Moore, 1985 A first New Zealand record . For the opportunity to examine material in their care ...... 76 we thank R. M. Emberson and J. M. W. Marris brevicollis Bates, 1874 E ...... 74 (Entomology Research Museum, Lincoln University, promptus (Erichson, 1842) A first New Zealand Lincoln), R. Didham (University of Canterbury, record ...... 77 Christchurch), P. M. Johns and S. D. Pollard (Canterbury puncticollis Bates, 1874 E ...... 75 Museum, Christchurch), B. H. Patrick (Otago Museum, Genus Haplanister Moore, 1996 A ...... 77 Dunedin), J. W. Early (Auckland Institute and Museum, crypticus Moore, 1996 A ...... 78 Auckland), R. L. Palma and P. Sirvid (Museum of New Zealand Te Papa Tongarewa, Wellington), B. I. P. Barratt Genus Kiwiharpalus new genus E ...... 79 (AgResearch, Mosgiel), P. Howe (Timaru), J. Nunn townsendi new species E ...... 79 (Dunedin), J. I. Townsend (Levin), S. Thorpe (Auckland), Genus Pholeodytes Britton, 1962 E ...... 80 T. A. Weir (Australian National Insect Collection, Canberra, cerberus Britton, 1964 E ...... 81 A.C.T.), O. Montreuil and T. Deuve (Muséum d’Histoire E helmorei new species ...... 83 Naturelle, Paris, France), C. Taylor and R. Booth (Natural E nunni new species ...... 82 History Museum, London, U.K.), R. Davidson and R. E palmai new species ...... 81 Ward (Carnegie Museum, Pittsburgh, Pennsylvania, townsendi Britton, 1962 E ...... 82 U.S.A.), M. Thayer and P. P. Parrillo (Field Museum of Natural History, Chicago, Illinois, U.S.A.), R. Poggi (Museo Civico di Storia Naturale, Genova, Italy). CONTENTS We thank B. P. Moore (Australian National Insect Acknowledgments ...... 13 Collection, Canberra) for his help in confirming the identity Introduction ...... 14 of adventive species and helpful advice in solving some Morphology and terminology ...... 17 nomenclatural problems. Methods and conventions ...... 21 We also wish to thank the following referees for their Taxonomic treatments ...... 24 helpful comments and suggestions for improving the Tribe Harpalini ...... 24 manuscript: H. Goulet (Agriculture and Agri-Food Canada, Key to subtribes of New Zealand Harpalini ...... 25 Ottawa), R. L. Palma (Museum of New Zealand Te Papa Alternative key to genera of New Zealand Harpalini Tongarewa, Wellington), R. J. B. Hoare and T. K. Crosby ...... 25 (Landcare Research, Auckland), and B. P. Moore (Australian Subtribe Anisodactylina ...... 26 National Insect Collection, Canberra). 14 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Thanks are also extended to B. Rhode (Landcare provide solid foundations for studies of other types, much Research, Auckland) for her help with distribution maps, in the same way as the work done by Lindroth between digital photographs and illustrations, and other technical 1961 and 1969 for Canada and Alaska, and Darlington assistance. between 1962 and 1968 for New Guinea. We are grateful to D. W. Helmore for the habitus and In addition to paper-based publications the authors frontispiece drawings. Finally, we are most obliged to T. publish the New Zealand Carabidae website (http:// K. Crosby (Landcare Research, Auckland) and O. R. W. www.landcareresearch.co.nz/) which maintains up-to-date Sutherland (former Science Manager, Landcare Research) information on New Zealand carabids, including digital for their encouragement and for allocating resources images, recent literature, as well as additions and corrections towards completion of this work. to previous publications. Most of the support for this research was provided through a subcontract of the Biosystematics of New Taxonomic history Zealand Terrestrial Invertebrates programme (Foundation for Research, Science and Technology Contract no There has been little work done on the faunistics of the C09X002). New Zealand Harpalini since the earliest descriptions of Hypharpax antarcticus and Triplosarus novaezelandiae by Laporte de Castelnau in 1867. No identification keys or INTRODUCTION taxonomic overview (except for the catalogue of Larochelle The tribe Harpalini belongs to the subfamily Harpalinae & Larivière 2001 and the checklist of Larochelle et al. 2004) (Coleoptera: Carabidae) which contains over 19 000 taxa. have been published until now, but keys including some Molecular sequence data indicate that Harpalinae radiated native taxa have been published by Sloane (1898 and 1920; in the Cretaceous Period (Ober 2002). Australian taxa), Noonan (1973; world Anisodactylina gen- The Harpalini form a diverse group, including over 240 era), Moore (1977; Australian taxa), and Matthews (1980; genera and subgenera, and approximately 2 000 species South Australian Carabidae genera). distributed in all biogeographic regions of the world. The Prior to this revision 13 genera and 36 species of present faunal review records 20 genera and 57 species for Harpalini were known from New Zealand. Following the New Zealand. This should constitute the near totality of work of Laporte de Castelnau (1867–1868), most the fauna. indigenous genera and species were described before 1920 Compared with New Zealand, the Australian Harpalini by Broun (1880–1914; 3 genera, 15 species) and Bates are more diverse with over 160 species distributed in about (1874, 1878; 3 genera, 6 species). Britton (1962, 1964a–b) 20 genera (Moore et al. 1987), but the fauna remains largely and Moore (1996) provided the most recent descriptions unrevised. for 2 genera, Pholeodytes and Haplanister respectively, The present work offers a concise faunal taxonomic and 6 species (including 2 in Parabaris and Syllectus). This revision of the New Zealand Harpalini, based on the study formed the bulk of the taxonomic work on New Zealand of adults contained in local and overseas collections. It Harpalini until now. No larval descriptions are yet available represents a first modern attempt to bring together the for this tribe. scattered information dealing with the group. If taxonomic progress has been slow until now, the collecting effort has been more intensive from the 1960s The goals of this revision are straightforward: to provide onward, so that New Zealand entomological collections an inventory of New Zealand taxa, a concise treatment of and museums are now replete with Harpalini material from their taxonomy, identification keys to genera and species, all areas of the country. For this reason, it seemed timely to and a summary of available information on species provide a taxonomic revision for this group, one that distribution, ecology, biology, and dispersal power. includes descriptions and keys that take into account this It is one step in the authors’ overall goal of attaining an new information. overall understanding of the carabid fauna within a The main taxonomic works that have contributed to reasonable time frame, and to make relatively large amounts advancing knowledge on world and New Zealand Harpalini of information available for practical use by a wide range of are: Sloane (1898, key to Australian genera); Jeannel (1942, end-users. The methodology involves less gamma taxonomy revision of France and world classification); Basilewsky but more intensive field work, and it is based on the concept (1950 and 1951, African revision); Lindroth (1968, revision of ‘practical taxonomy’ described by Darlington (1971) of Canada, Alaska, and northern half of U.S.A.), Darlington which aims to provide “a floor plan for more detailed (1968, revision of New Guinea); Habu (1973, revision of taxonomic, ecological, zoogeographical, and evolutionary Japan); Noonan (1973, generic revision and classification studies.” of world Anisodactylina; and 1976, world catalogue of It is hoped that this kind of faunal taxonomy will supraspecific taxa of Harpalini); Goulet (1974, revision of Fauna of New Zealand 53 15 genus Pelmatellus); Moore (1977, key to Australian Subtribe Harpalina. Representatives of this group occur subtribes); Matthews (1980, key to South Australian in all zoogeographical regions, mostly in tropical and tem- genera); Moore et al. (1987, Australian catalogue); Bousquet perate areas. Approximately 60 genera are known. The & Larochelle (1993, Nearctic catalogue); Serrano et al. (1994, taxa occurring in New Zealand were introduced from Aus- karyotypical study); Ball & Bousquet (2001, key to tralia and the Holarctic. supraspecific taxa, North America); Larochelle & Larivière Subtribe Pelmatellina. Members of this small group (2001, catalogue of New Zealand Carabidae); Kataev exhibit a Gondwanan distribution pattern in Australia, New (2002a, new genus of Australian Anisodactylina); Löbl & Smetana (2003, Palaearctic catalogue). Zealand, Andean South America, and Central America, with some taxa reaching the southwestern U.S.A. About 8 gen- era were described before this revision. Higher classification Pelmatellina are considered the sister group of The monophyly of the subfamily Harpalinae, to which Anisodactylina based on the shared spongily pubescent belongs the tribe Harpalini, has recently been supported male protarsi (Noonan 1973; Goulet 1974). The current by molecular sequence data (Ober 2002) and larval mor- study on New Zealand taxa also agrees with Noonan (1976) phology (Arndt 1998). on the character of the penultimate segment of the labial According to Bousquet & Larochelle (1993) the palpi which is bisetose (most genera) or trisetose taxonomic limits of the tribe Harpalini are fairly stable (Kupeharpalus new genus) in Pelmatellina; not strictly although the monophyly of this taxon remains to be tested. bisetose as suggested by Goulet (1974). Both Noonan (1973, The main contributor to the higher classification of this 1976) and Goulet (1974) indicated that pelmatelline genera group was Noonan (1973, 1976) who studied the differ from anisodactyline genera by the glabrous apex of taxonomy, phylogeny, and zoogeography of the subtribe the prosternal lobe. Four pelmatelline genera are now Anisodactylina and provided a synopsis of supra-specific known from New Zealand, three of which share this taxa of the tribe Harpalini. character (Lecanomerus, Syllectus, and Hakaharpalus new The supraspecific classification proposed by Noonan, genus). Kupeharpalus new genus provides the exception and based on the earlier work of van Emden (1953), to this rule in having a prosternal lobe apically pubescent grouping genera into 4 subtribes (Anisodactylina, but in other respects sharing the characters of Lecanomerus. Harpalina, Pelmatellina, and Stenolophina), is generally Further elucidation of character evolution in the Pelmatellina accepted worldwide although somewhat difficult to apply in certain cases (e.g., taxonomic limits of Pelmatellina). will have to wait until all subtribes of Harpalini are revised This classification is followed here. The subtribes Harpalina, on a world basis. Pelmatellina, and Stenolophina still need an analysis such Subtribe Stenolophina. Most species of this subtribe as provided by Noonan (1973) for Anisodactylina. occur in the warm temperate and tropical regions, with 35 Subtribe Anisodactylina. Members of this group are dis- genera or so recorded worldwide. The morphology of New tributed worldwide. About 40 genera are known (Kataev Zealand stenolophine genera, including Kiwiharpalus new 2002a) from two genus-groups (Notiobii and Anisodactyli). genus, is consistent with the diagnostic characters pro- According to Ball & Bousquet (2001), the Notiobii are vided by Noonan (1976) for this subtribe. principally in the Southern Hemisphere, showing a Noonan (1976) recorded two small endemic genera in Gondwanan distribution pattern, whereas the Anisodactyli the Australian Region (Euthenarus and Pholeodytes), to occur mostly in the Afrotropical and Holarctic Regions. which the current revision adds the new genus Most New Zealand genera have the Notiobii character of Kiwiharpalus. Noonan also indicated that species of several the complete transverse suture between mentum and other genera occurring in the Australian Region may be submentum. This represents the plesiomorphic state in primarily centred in the Oriental Region and spreading Anisodactylina. Only the endemic Gaioxenus has the trans- only to the outer limit of the Australian Region or are verse suture laterally incomplete (mentum and submentum Australian-centred taxa that may have originated from fused only medially). This is usually regarded as a charac- Oriental stock. ter of the Anisodactyli, but exceptions have been observed Ball & Bousquet (2001) placed the North American by Noonan (1976) in other Southern Hemisphere taxa, stenolophine genera into 2 genus-groups, Stenolophi and e.g., within species of Anisostichus and subgenera of the Polpochili. According to the literature, one important Notiobia lineage, and may represent examples of parallel character defining the genus-group Stenolophi is the evolution. Noonan (1973) believed that the subtribe ventrally pubescent male protarsi as opposed to the absence Anisodactylina forms a monophyletic group but he was of such pubescence in Polpochili. The study of this unable to state that the group is defined on the basis of character in taxa indigenous to New Zealand suggests that clearly apomorphic character states. species of Pholeodytes (endemic) and Euthenarus (not 16 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) endemic) could belong to the Stenolophi. This character latitarsis), 2 natives in common with the South Island could not be studied in Kiwiharpalus which is known only (Hypharpax antarcticus, Lecanomerus latimanus), and one from females. However, an Australian revision and a world shared with the North and South Islands (Euthenarus reclassification and phylogeny of supra-specific taxa of puncticollis), and 3 adventives. Harpalini are so far unknown Stenolophina are needed in order to uncover the true from New Zealand’s subantarctic islands. evolutionary history of this subtribe. A total of fourteen (14) adventive species (about 25% of Harpalini) occur throughout New Zealand, mostly in the North Island (Map 7; especially in WN, ND, WI). The Geographic distribution and faunal composition majority of adventive species probably originated from The level of endemism of the New Zealand Harpalini is Australia apart from 2 Harpalus species and Anisodactylus 75% at the species level (42 out of 57 species) and 55% at binotatus (from the Palearctic Region), and Haplanister the generic level (11 out of 20 genera). The indigenous crypticus (of unknown origin). genera Hypharpax, Lecanomerus, and Euthenarus have The areas of New Zealand so far known to contain the representatives in Australia. The genera Anisodactylus, highest diversity (Map 4) are: NN (23 species), ND (21 Gnathaphanus, Notiobia, Harpalus, Egadroma, and species), WN (17 species). The areas with the greatest Haplanister are adventive. number of New Zealand endemics (Map 5) are: NN (16), The overall distribution of New Zealand Harpalini is ND (13), BP (11). summarised in Table 1. Some Harpalini are restricted to a single area (Map 6). Species distributions are clearly defined and largely Currently, the areas with such species are: NN (9), ND (6), allopatric. Even species of a single genus, occurring in the TH (4), BR (1), BP (1), CL (1), MC (1), CH (1). The same general areas of New Zealand are mostly allopatric South Island northwest (NN, BR) and the far north of the within these areas (e.g., Tuiharpalus, TH–ND; New Zealand (ND, TH) appear to have been the reservoirs, Kupeharpalus, ND; Pholeodytes, NN; Hakaharpalus BR– in geological time, of much of the genetic diversity in New NN–SD). Zealand Harpalini, with several species currently restricted Three genera (Gaioxenus, 1 species; Parabaris, 3 to these regions. This trend is reflected at the generic level species; Kupeharpalus, 3 species) are confined to the North with Hakaharpalus occurring only in BR–NN–SD, Island. The genus Allocinopus (7 native species) occurs Pholeodytes in NN, Kupeharpalus in ND, Maoriharpalus mostly on the North Island, except for 2 species, A. and Kiwiharpalus in TH, and Tuiharpalus in TH–ND. sculpticollis which is also found on the South Island, and A. Table 2 shows the genera and species shared with latitarsis which is endemic to the Chatham Islands (CH). Australia, New Caledonia, Norfolk Island, and Lord Howe Two genera (Hakaharpalus, 5 species; Pholeodytes, 5 cave- Island. Ten (10) species shared with these regions are dwelling species) are found only on the South Island and adventive in New Zealand. Three indigenous genera are restricted to the NN–SD region. Two genera are restricted (Hypharpax, Lecanomerus, Euthenarus) are shared with to the Three Kings Islands (TH): Maoriharpalus (1 species) Australia (eastern Australian mainland and Tasmania). and Kiwiharpalus (1 species). There is no genus endemic to the Chatham Islands (CH). Thirty-five (35) Harpalini species occur on the North Ecology, biology and dispersal power Island, with 16 native species restricted to it. Thirty-one No formal detailed study of the natural history of indi- (31) species are distributed on the South Island, with 14 vidual species of New Zealand Harpalini has ever been native species restricted to it. Only 4 indigenous species conducted although Larochelle & Larivière (2001) summa- are shared between these two main islands (Allocinopus rised information available from the literature, material in sculpticollis, Triplosarus novaezelandiae, Syllectus entomological collections, personal communications from anomalus, and Euthenarus puncticollis). Stewart Island carabid collectors, and their own personal field observa- has no endemic taxa, but shares 2 indigenous species: tions. Triplosarus novaezelandiae (with North Island and South Native species are mostly subapterous and live within Island), Euthenarus brevicollis (with South Island). Six (6) the confines of native habitats, mostly forests (especially species occur on the Three Kings Islands (TH), including 4 along streams) and wet habitats, also tussock grasslands endemics (Maoriharpalus sutherlandi, Tuiharpalus crosbyi, and caves (2 Syllectus, 5 Pholeodytes). The cave-dwelling T. gourlayi, Kiwiharpalus townsendi), 1 adventive, and 1 species are all troglobitic, except Syllectus magnus which indigenous species in common with the North Island is troglophilous, occurring at the entrance of caves. Most (Lecanomerus sharpi). Seven (7) species occur on the Harpalini species are hygrophilous (moisture-loving) living Chatham Islands (CH), including 1 endemic (Allocinopus at the surface of the soil and in leaf litter, also in caves Fauna of New Zealand 53 17

(Syllectus, Pholeodytes), and occasionally on plants and The Department of Conservation’s Species Ranking trees. Two native species are typically found along coastal System is summarised in Table 3. lowlands: Triplosarus novaezelandiae (on beaches and sand dunes), Allocinopus belli (coastal forests). Dispersal in When the above criteria are applied, new knowledge native species is achieved by running over the ground; brought forward in the present revision suggests that 24 most species are moderate runners, except for the long- endemic species of Harpalini (over 50% of native species) legged, fast-running cave species (Syllectus, Pholeodytes). known from 10 populations or fewer may be of potential In general Harpalini have relatively short legs and, conservation concern. sometimes, strongly reduced eyes which are indicative of All but two of these species are new to science and all strong burrowing habits. species are taxonomically highly distinctive, have limited All adventive species are macropterous and live mostly dispersal power, are often geographically localised in in highly modified environments (often around human threatened habitats, and represented in collections by dwellings), except for Haplanister crypticus which has relatively few specimens collected over many decades, which managed to invade native forests. may indicate rare or highly specialised species. The collecting period of teneral individuals suggests These species of special interest are: Allocinopus belli that Harpalini species may be either spring-breeders or new species, A. bousqueti new species, A. wardi new summer-breeders. For most species adults are active during species, Maoriharpalus sutherlandi new species, Parabaris all months of the year, but are generally less active during hoarei new species, P. lesagei new species, Tuiharpalus cooler months. clunieae new species, T. crosbyi new species, T. gourlayi There are no data available on the feeding preferences (Britton), T. hallae new species, T. moorei new species of Harpalini native to New Zealand. Larochelle (1990), in (Anisodactylina); Hakaharpalus cavelli (Broun), H. his review of food preferences of the Carabidae of the davidsoni new species, H. maddisoni new species, H. world, showed representatives of this tribe to be patricki new species, H. rhodeae new species, omnivorous, mostly phytophagous species. Ecomorph- Kupeharpalus embersoni new species, K. johnsi new ological adaptations providing further evidence for this species, Lecanomerus marrisi new species, Syllectus gouleti feeding-type in adults and larvae have been documented new species (Pelmatellina); Kiwiharpalus townsendi new by Sharova (1960, 1981), Acorn & Ball (1991), and Zetto species, Pholeodytes cerberus Britton, P. helmorei new Brandmayr et al. (1998). The mandibles of Hakaharpalus, species, P. nunni new species, and P. palmai new species Kiwiharpalus, Syllectus, Pholeodytes, and Maoriharpalus (Stenolophina). are unusually long among native Harpalini, which may suggest a specialised type of feeding. In addition, the strongly emarginate labrum of Maoriharpalus is MORPHOLOGY AND TERMINOLOGY reminiscent of, although not necessarily equivalent to, the The main diagnostic features of Harpalini are: body usu- condition observed in Licinini which feed on hard-bodied ally rather stout, with relatively short appendages; head invertebrates, e.g., snails. with a single pair of supraorbital setiferous punctures; mandibles usually relatively short, without setae in scrobes; Conservation status posterior angles of pronotum usually without a setiferous The Department of Conservation has responsibility for puncture; elytral apex neither truncate nor crossed subapi- protecting and conserving New Zealand’s native plants cally; median lobe of aedeagus with basal bulb well devel- and animals. The Department’s Species Priority Ranking oped in most taxa, shaft usually strongly arcuate; parameres System established by Molloy et al. (1994) provides crite- of aedeagus usually short and broad, conchoid (shell-like) ria for scoring species according to various levels of threat, or ovate, similar in shape with right paramere usually so that management and/or recovery plans can be subse- smaller. quently established. A list of priority invertebrate species A more detailed description of the tribe based on New for conservation was established in this way by Molloy et Zealand representatives is available on page 24. Figures 1– al. (1994). McGuinness (2001) developed species profiles 31 provide a basic understanding of the morphological for species on the list, providing additional descriptive structures used to describe and identify Harpalini genera information to initiate or support key conservation ac- and species. A glossary of technical terms is also provided tions. In addition, McGuinness (2001) added a number of (Appendix A, p. 89). invertebrates of potential conservation interest to the origi- nal list. No Harpalini species has been listed in these docu- ments. 18 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Table 1. Summarised distributions for New Zealand Harpalini. Two-letter area codes follow Crosby et al. (1976, 1998). A = Adventive taxa. Species North Island ND A K C L WO BP TK T O HB GB RI W I WN WA Anisodactylina Allocinopus angustulus HB RI WI WN W A A. belli BP A. bousqueti BP GB A. latitarsis A. sculpticollis BP TK TO HB GB RI WI WN W A A. smithi AK WO BP TK TO A. wardi CL Anisodactylus binotatusA WN Gaioxenus pilipalpis CL WO BP TK TO RI WI WN Gnathaphanus melbournensisA HB Hypharpax antarcticus H. australisA ND AK CL WO BP TK TO HB GBRI WI WN Maoriharpalus sutherlandi Notiobia quadricollisA ND Parabaris atratus ND A K CL WO BP TK GB P. hoarei ND P. lesagei BP TK WN Triplosarus novaezelandiae ND A K CL WO BP HB WI WN Tuiharpalus clunieae ND T. crosbyi T. gourlayi T. hallae ND T. moorei ND Harpalina Harpalus affinisA RI WI WN W A H. australasiaeA BP HB GB WI WN W A H. tardusA Pelmatellina Hakaharpalus davidsoni H. maddisoni H. patricki H. rhodeae Kupeharpalus barrattae ND A K CL K. embersoni ND K. johnsi ND A K CL BP Lecanomerus atricepsA ND A K CL BP WIWN L. insignitus ND L. latimanus L. marrisi L. obesulus L. sharpi ND AK CL WO BP TK TO HB GBRI WI WN WA L. verticalisA ND A K CL WO BP GBWIWN L. vestigialisA ND A K CL WO BP HBGBWIWN Syllectus anomalus ND AK CL WO BP TK TO HB GBRI WI WN WA S. gouleti S. magnus Stenolophina Egadroma piceaA ND A K CL Euthenarus bicolorA ND A K CL WO BP TO HB GB E. brevicollis E. promptusA TO HB WI WN W A E. puncticollis ND AK CL WO TK TO HB GBRI WI WNWA Haplanister crypticusA ND AK CL WO BP TK TO HB GBRI WI WN WA Kiwiharpalus townsendi Pholeodytes cerberus P. helmorei P. nunni P. palmai P. townsendi Fauna of New Zealand 53 19

Table 1 (continued)

Offshore Species South Island Islands SD NN M B K A BR WD NC M C S C M K O L C O DN FD S L SI Anisodactylina Allocinopus angustulus A. belli A. bousqueti A. latitarsis CH A. sculpticollis SD NN BR WD A. smithi A. wardi Anisodactylus binotatusA MC SC OL CO DN Gaioxenus pilipalpis Gnathaphanus melbournensisA KA NC MC Hypharpax antarcticus NN MB K A NC MC SC MK OL CO DN CH H. australisA SD NN MB K A MC SC MK OL CO CH TH Maoriharpalus sutherlandi TH Notiobia quadricollisA Parabaris atratus P. hoarei P. lesagei Triplosarus novaezelandiae SD NN WD MC DN S L SI Tuiharpalus clunieae T. crosbyi TH T. gourlayi TH T. hallae T. moorei Harpalina Harpalus affinisA H. australasiaeA MC H. tardusA MC Pelmatellina Hakaharpalus davidsoni NN H. maddisoni NN H. patricki NN H. rhodeae SD NN Kupeharpalus barrattae K. embersoni K. johnsi Lecanomerus atricepsA SD NN MC L. insignitus L. latimanus KA MCSC CO DN S L CH L. marrisi MC L. obesulus SD NN MB BR WD NC MC OL CO FD SL L. sharpi TH L. verticalisA NN L. vestigialisA NN MCSC DN CH Syllectus anomalus SD NN MB K A BR WD NC MC DN FD S L S. gouleti BR S. magnus NN BR Stenolophina Egadroma piceaA Euthenarus bicolorA NN E. brevicollis K A NC MC SC MK OL CO DN FD S L SI E. promptusA SD NN MB K A BR MC E. puncticollis SD NN BR WD CH Haplanister crypticusA NN MCSC CH Kiwiharpalus townsendi TH Pholeodytes cerberus NN P. helmorei NN P. nunni NN P. palmai NN P. townsendi NN 20 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Table 2. Taxa shared with Australia, New Caledonia, Norfolk Island, and Lord Howe Island. X = present; [ ] = adventive; — = absent.

Species New Australia Tasmania New Norfolk Lord (macropterous) Zealand (mainland) Caledonia Island Howe Island

Anisodactylina Gnathaphahus [X] X X X [X] [X] Gnathaphanus melbournensis [X] X X —— [X] Hypharpax X X X —— [X] Hypharpax australis [X] X X —— [X] Notiobia [X] X X X [X] [X] Notiobia quadricollis [X] X X —— — Harpalina Harpalus [X] X — —— — Harpalus australasiae [X] X X —— — Pelmatellina Lecanomerus XX X X—— Lecanomerus atriceps [X] X — —— — L. verticalis [X] X X —— — L. vestigialis [X] X X —— — Stenolophina Egadroma [X] X X X [X] [X] Egadroma picea [X] X X X — — Euthenarus X X X — [X] — Euthenarus bicolor [X] X — — [X] — E. promptus [X] X X — [X] —

Table 3. Department of Conservation Species Priority Invertebrates are then grouped into 3 categories depend- Ranking System (Molloy & Davis, 1994; McGuinness, ing on the score they receive from the ranking system. 2001). Designed to categorise threatened species A: Highest priority threatened species for conservation according to their urgency for conservation. (bold = action. criteria more readily applicable to Harpalini based B: Second priority threatened species for conservation on current taxonomic and biological knowledge) action. Plants and animals are scored using 5 factors, encom- C: Third priority threatened species for conservation ac- passing 17 criteria. tion. 1. Distinctiveness: taxonomic distinctiveness. In addition, 4 other specialist categories are used: 2. Status: number of populations; mean population X: Species that have not been sighted for a number of size; size of largest population; geographic distri- years and are presumed extinct. bution; condition of largest population; and the popu- lation decline rate. I: Species about which little is known, but based on exist- ing knowledge are considered to be under threat. 3. Threats: legal protection of habitat; habitat loss rate; predators/harvest impact; competition; and other fac- O: Species that are threatened in New Zealand but are tors affecting survival. known to be secure in parts of their range outside New Zealand (no invertebrate so far listed in this 4. Vulnerability: habitat and/or diet specificity; reproduc- category). tive and/or behavioural specialisation; and cultivation/ captive breeding potential. M: Species that are [apparently] rare or localised, and of cultural importance to Maori. 5. Values: Maori cultural values; Pakeha cultural values. Fauna of New Zealand 53 21

METHODS AND CONVENTIONS vegetation into water, digging at the base of plants (e.g., Lupinus), pouring water over ground, treading soil with Materials the feet, sweeping the vegetation, using Malaise traps, This revision is based on 12 years of extensive field work collecting with a head lamp or torch (e.g., in caves; on trees carried out by the authors in over 500 localities, a survey at night), light trapping (especially for adventive species), of the literature up to May 2004, and the recording of sifting garden compost, and turning drift shore material. information associated with over 5350 adult specimens Pitfall trapping, especially in forests (along streams) and from the following entomological museums and collections: in coastal dunes is the most reliable method for assessing AMNZ Auckland Institute and Museum, Auckland, the presence, community composition, and locomotory New Zealand. activity of harpalines. ANIC Australian National Insect Collection, Canberra, Adults are best preserved dry. All life stages can be Australia. collected in 70–75% ethanol. If a molecular study is BBNZ B. I. P. Barratt private collection, Dunedin, New intended, adults as well as immatures can be kept in 95– Zealand. 100% ethanol. BMNH The Natural History Museum, London, U.K. All specimens should be labelled with the locality CMNH Carnegie Museum of Natural History, Pitts- (including area code: Crosby et al. 1976, 1998, and burgh, Pennsylvania, U.S.A. geographical coordinates such as latitude and longitude), CMNZ Canterbury Museum, Christchurch, New Zea- collection date, collector’s name, and biological data (e.g., land. general habitat, microhabitat, behaviour). FMNH Field Museum of Natural History, Chicago, Il- Most features of the external morphology and the male linois, U.S.A. genitalia can be viewed under an ordinary dissecting ITNZ J. I. Townsend private collection, Levin, New microscope. It is necessary to relax and dissect male Zealand. specimens in order to study their genitalia. JNNZ J. Nunn private collection, Dunedin, New Zea- Male genitalia can be dissected as follows. Pinned land. specimens (individually or in batches) are warmed for 5– LUNZ Entomology Research Museum, Lincoln Uni- 10 minutes in hot alcohol (70–75% ethanol). Once softened, versity, Lincoln, New Zealand. each specimen is transferred to a cavity slide containing MCSN Musei Civico di Storia Naturale, Genova, Italy. ethanol. A pair of fine forceps is used to extract the male MNHN Muséum National d’Histoire Naturelle, Paris, genitalia from the abdomen. This is done under the France. microscope by inserting the forceps into the anus, cutting MONZ Museum of New Zealand Te Papa Tongarewa, through the lateral membranes that unite the last two tergites Wellington, New Zealand. and ventrites, pulling out the aedeagus and associated genital NZAC New Zealand Arthropod Collection, Landcare ring, separating these structures from each other, and Research, Auckland, New Zealand. cleaning the aedeagus from any residues and detaching the OMNZ Otago Museum, Dunedin, New Zealand parameres. The dissected genitalic structures are then PHNZ P. Howe private collection, Timaru, New Zea- transferred to a new cavity slide containing glycerol for land. further study. After examination, the male genitalia are UCNZ Department of Zoology, University of Canter- mounted on cards or points and re-attached to original bury, Christchurch, New Zealand. specimens for permanent storage. Specimen-based information from NZAC is being databased and will be made available online on the NZAC Revision process NZBUGS website (http://www.landcareresearch.co.nz/). The main steps followed in the course of this study are listed here with the hope that this will help future students Collecting and preparation of Carabidae: 1. Borrowing of adult specimens from all available ento- Adults of Harpalini are generally collected by hand by mological collections (private and institutional). turning ground debris. However, special techniques are used to collect large series or population samples for quan- 2. Labelling of borrowed specimens with the acronyms of titative studies. These include (in order of decreasing us- the lending collections. age): pitfall trapping, turning logs and stones, raking or 3. Grouping of specimens based on overall similarities and sifting the leaf litter (especially for small species), treading differences in external morphology. 22 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

4. Grouping of recognised morphological units by areas of must allow some degree of variation when working with New Zealand (area codes of Crosby et al., 1976 & individual specimens. 1998) from North to South, West to East, and by off- The male genitalia offer the most stable characters shore islands. This facilitates the evaluation of struc- and the ultimate criteria for species recognition. The tural variation between and within populations across second best diagnostic character for the majority of taxa is the geographic range of each putative species. the configuration of the pronotum. 5. Dissection of male genitalia from at least 5 populations Although it is necessary to fully dissect male genitalia per area. Additional specimens sometimes need to be within the context of taxonomic revisions, it is often enough dissected from some areas (e.g., from WA, WN, SD, to partially pull out the apex of the aedeagus with a pin at NN) where a high degree of variation may be observed the time of mounting specimens in order to see enough of between and within populations. About 650 male speci- the genitalia for identification. mens were dissected in the course of this study. Parameres of the aedeagus were found to vary little between species and were not illustrated. In the Harpalini, 6. Identification of putative species based on male genitalia the internal sac of the male aedeagus is either armed and drawing of their genitalic features (lateral and dor- (provided with scales, teeth, or spines) or unarmed. These sal views of aedeagus). conditions have been stated although not illustrated for 7. Correlation of results from the genitalia study with char- each species. Illustrating these characters in detail would acters of the external morphology at the species level have required the eversion of the internal sac, which was (corroborated, when possible, by geographical and bio- beyond the scope of this revision. The female genitalia logical information). were not studied either. There was sufficient diagnostic 8. Photography of pronotum and whole insect for each information provided by other characters. species. 9. Description of taxa based on a character list developed Identification keys from previously published works, the study of popu- lation samples in steps 6 and 7, and the drafting of a Keys are somewhat artificial. They are intended as an aid description for one species (often the type species) to identification, not a statement of the authors’ opinion from each described genus. This involves the descrip- on phylogenetic relations. Additional supporting charac- tion of each species in detail followed by the transfer ters (e.g., distribution) have sometimes been included be- of selected characters from the species descriptions to tween key couplets to help identification. the generic descriptions. 10. Comparing circumscribed species against the types of Illustrations and digital photographs already described species and application of existing Illustrations (except habitus drawings and Fig. 114–225), names or new names. including maps, were prepared from pencilled drafts that 11. Preparing identification keys from descriptions. were digitised, finished, and laid out using the software package CorelDraw graphics suite. Colour photographs of whole insects and pronota were captured through a Leica Taxonomically relevant characters MZ-12 stereomicroscope, a 3CCD video camera, a The characters presented in the descriptions are subsets of LeicaDC500 digital camera, and the increased-depth-of- the totality of adult characters (about 100) studied, and field computer system Auto-Montage (Synoptics U.K.). represent the most important differences between, or vari- Further photo-processing was done with the software pack- ation amongst, closely related taxa. Characters or states of ages PhotoShop and CorelDRAW graphics suite. characters not mentioned in the species descriptions are as described in generic descriptions. Body length was measured from apex of mandibles to Generic concept apex of elytra (with the specimen in dorsal view), and is A genus should be a monophyletic group composed of one cited as a range. or more species separated from other genera by a decided Characters with the highest diagnostic value at the gap. The phylogenetic framework to study Australasian species level have been photographed or illustrated, Harpalini, however, is insufficiently elaborated to test this including the most diagnostic aspects of the male genitalia. hypothesis for New Zealand genera. Consequently, exist- Most illustrations provided in this work represent the ing generic concepts have in general been accepted. In addi- most commonly encountered state of a character. The user tion, new genera are proposed for species not fitting the Fauna of New Zealand 53 23 correlated character complex of species included in already Biostatus described genera. Recognition of these generic taxa pro- This is indicated for all genera and species (A=Adventive; vides new hypotheses that will hopefully be tested by E=Endemic; N=Native, not endemic). The biostatus cat- future students of the higher classification of Harpalini; egories used are defined in the Glossary (Appendix A, p. this must be done on a world basis or at least in an Aus- 89). A combination of criteria was used to assess whether tralasian context. taxa were adventive including: recency of first New Zea- A cladistic analysis, preferably integrating land record in the literature and collections; fit of current morphological and genetic information, is needed to geographical and ecological distribution with recognised determine the phylogenetic position of New Zealand genera natural patterns, or similarity of such distribution with within the Harpalini. Only then can an attempt be made to that of other adventive arthropods; and dispersal ability, decipher the evolutionary history of the New Zealand taxa, especially in relation to flightlessness and distance from e.g., to confirm or reject the hypothesis that certain genera the nearest overseas populations. are Gondwana relicts, to reconstruct the sequence of speciation and colonisation events, and to understand their evolution in general or that of their habitat relationships. Type data The primary types of native species were examined. Such information is listed in the following format: type status Species concept (holotype, lectotype, etc.) followed by sex, acronym of The species concept used here is biological, inferred from entomological collection or museum serving as repository, morphological characters (especially male genitalia) hy- and original label data with a forward slash (/) indicating a pothesised to constitute barriers to interbreeding and hence different label. Only type localities are provided for to gene flow between the different species. This is cor- adventive species. roborated, when possible, by geographic and biological information, but is not tested by genetic or ethological Geographic distribution investigations. This species concept requires the assump- For New Zealand distribution records, the area codes of tion that reproductive (genetic) continuity or isolation Crosby et al. (1976, 1998) are given in alphabetical order among natural populations is evidenced by continuity or for the North Island, South Island, Stewart Island, and the discontinuity in characters of external morphology and Offshore Islands, respectively. When appropriate, the genital structures provided by the study of population extralimital distribution (outside New Zealand and its off- samples. shore islands) is also included, as well as first New Zealand As generally observed in Carabidae, the most important records of adventive species. Full distributional informa- characters to discriminate Harpalini species are the male tion is given for species known from ten (10) localities or genital structures, particularly the aedeagus. In the majority fewer. Appendix B (p. 91) contains a list of the main col- of New Zealand genera, many external characters are found lecting localities and their geographic coordinates. to vary within species, or the range of their variation Two-letter abbreviations for the area codes of Crosby overlaps with that of closely related species, and similarities et al. (1976, 1998) used in this publication are as follows or differences in external morphology are not always (see Maps 1-3): congruent with the study of genitalia. Accurate species New Zealand. North Island: AK, Auckland; BP, Bay identification is generally impossible without an of Plenty; CL, Coromandel; GB, Gisborne; HB, Hawke’s examination of male aedeagus. Therefore, in most cases, Bay; ND, Northland; RI, Rangitikei; TK, Taranaki; TO, females can only be reliably identified by association with Taupo; WA, Wairarapa; WI, Wanganui; WN, Wellington; males. Fortuitously, identification is facilitated by the fact WO, Waikato. South Island: BR, Buller; CO, Central that New Zealand species are largely allopatric. Otago; DN, Dunedin; FD, Fiordland; KA, Kaikoura; MC, Mid Canterbury; MK, Mackenzie; NC, North Canterbury; NN, Nelson; OL, Otago Lakes; SC, South Canterbury; SD, Taxonomic arrangement Marlborough Sounds; SL, Southland; WD, Westland. Further study of Australasian Harpalini is needed before Stewart Island, SI. Offshore Islands: AN, Antipodes phylogenetic relationships can be hypothesised. In this Islands; AU, Auckland Islands; BO, Bounty Islands; CA, monograph, subtribes and genera are treated alphabetically Campbell Island; CH, Chatham Islands; KE, Kermadec while species are arranged according to their similarity in Islands; SN, Snares Islands; TH, Three Kings Islands. male genitalia and external morphology, which may or may Maps summarising species distributions by areas of not be indicative of phylogenetic relationships. New Zealand are provided on pp. 147-153. 24 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Material examined TAXONOMIC TREATMENTS This indicates the number of specimens examined and the Tribe HARPALINI acronym of their repositories. Diagnosis (New Zealand). Head with a single pair of supraorbital setiferous punctures. Body usually rather Ecology, biology, and dispersal power stout, with relatively short appendages. Mandibles usu- ally relatively short. Posterior angles of pronotum without The information provided is based on specimen label data, a setiferous puncture. Elytral apex neither truncate nor field observations from the authors, and the literature. In crossed subapically. order to eliminate spurious records an effort was made to summarise available information by using the smallest com- Description (New Zealand). Body length: 3.0–20 mm. Mostly pigmented and dark in colour, rarely depigmented mon denominator amongst the greatest number of observa- and testaceous. Generally glabrous and smooth. Body not tions for each species. The terminology and style of pres- pedunculate, usually rather stout, with relatively short entation adopted here follows Larochelle & Larivière appendages. Head with a single pair of supraorbital (2001). Most technical terms are also defined in the glos- setiferous punctures. Labrum usually transverse; apex sary (Appendix A, p. 89). straight or slightly emarginate medially, rarely strongly emarginate (Maoriharpalus); anterior margin with 6 References setiferous punctures. Clypeus narrower than distance be- tween antennal scapes; apex straight or slightly emarginate Under References, only the most important references are medially; each outer distal angle with one setiferous punc- given for each taxon, with an indication of their contents ture. Mandibles usually relatively short; scrobe without a between parentheses. setiferous puncture. Palpi visibly pubescent, rarely subglabrous; penultimate segment of labial palpi either plurisetose (with 4 setae or more), trisetose (with 3 setae), or bisetose (with 2 setae) on anterior margin. Antennae usually moderately long, reaching pronotal base; pubes- cence starting generally on antennomere 3, rarely on antennomere 2. Mentum generally with a tooth medially, moderately shorter than lateral lobes. Mentum and submentum usually separated by complete transverse su- ture. Each pronotal side with a setiferous puncture before middle. Posterior angles of pronotum each without a setiferous puncture. Scutellar striole of elytra usually present, inserted between interneurs 1 and 2. Procoxal cavi- ties uniperforated. Metepimeron visible as a lobe between metepisternum and ventrite 1. Elytra with apex rounded or angulate, not truncate; epipleura simple, not crossed sub- apically. Abdominal apex hidden from above. Male protarsi and usually mesotarsi laterally dilated and clothed with adhesive setae ventrally; male tarsi either spongily pubes- cent, biseriately pubescent, or rarely unmodified (i.e., sim- ple as in the female). Aedeagus (i.e., penis, median lobe) usually arcuate in lateral view, either asymmetrical (with ostium deflected laterally) or symmetrical (with ostium dorsal, not deflected laterally) in apical half in dorsal view; basal bulb well developed, feebly elbowed. Internal sac with or without armature (scales, teeth, or spines). Parameres short and wide, conchoid or ovate, slightly dif- ferent in shape, the right paramere being smaller (i.e., al- most as long, slightly narrower). Remarks. Klimaszewski & Watt (1997) provided a key to the subfamilies and tribes of Carabidae occurring in New Zealand. Fauna of New Zealand 53 25

Key to subtribes of New Zealand Harpalini (mostly setiferous micropores dorsally. Eyes moderately large, based on males) convex (Fig. 89). Tempora not inflated (Fig. 89) ...... (p. 36)... Gnathaphanus Macleay 1 Penultimate segment of labial palpi plurisetose (with 4 3(1) Mentum without a tooth medially (Fig. 18) ...... 4 setae or more; Fig. 9) on anterior margin ...... 2 — Mentum with a tooth medially (Fig. 14) ...... 6 — Penultimate segment of labial palpi trisetose (with 3 4(3) Eyes reaching buccal fissure ventrally (Fig. 21). Frons setae; Fig. 10) or bisetose (with 2 setae; Fig. 11) on with clypeo-ocular prolongations (Fig. 109). Body anterior margin ...... 3 length 6.5 mm or less ...... 2(1) Male protarsi biseriately pubescent (with 2 rows of ...... (p. 72)... Egadroma Motschulsky scale-like setae) ventrally (Fig. 13). Aedeagus — Eyes separated from buccal fissure ventrally (by 1–2x asymmetrical, with ostium strongly deflected to the maximum width of antennal scape) (Fig. 19). Frons left (Fig. 55) ...... (p. 50)... Harpalina without clypeo-ocular prolongations (Fig. 91). Body — Male protarsi spongily pubescent ventrally (Fig. 12). length 10 mm or more...... 5 Aedeagus asymmetrical (with ostium deflected to the 5(4) Mandibles (Fig. 91) and antennal scapes (Fig. 182) right (Fig. 35) or twisted (Fig. 39)) or symmetrical very long, about 6× their maximum width. Labrum (with ostium dorsal, not deflected laterally (Fig. 32)) strongly emarginate apically (Fig. 91). Mentum and ...... (p. 26)... Anisodactylina (in part) submentum separated by transverse suture (Fig. 22). 3(1) Penultimate segment of labial palpi trisetose on Pronotum suborbicular (Fig. 126). [TH] ...... anterior margin (Fig. 10) ...... 4 ...... (p. 40)... Maoriharpalus new genus — Penultimate segment of labial palpi bisetose on anterior — Mandibles (Fig. 87) and antennal scapes (Fig. 177) margin (Fig. 11) ...... 5 much shorter . Labrum straight or slightly emarginate 4(3) Frons without clypeo-ocular prolongations (Fig. 85) apically (Fig. 87). Mentum and submentum fused, not ...... (p. 26)... Anisodactylina (in part) separated by transverse suture (Fig. 24). Pronotum — Frons with clypeo-ocular prolongations (Fig. 107) .... rectangular (Fig. 121). [South Island and southern North ...... (p. 54)... Pelmatellina (in part) Island]...... (p. 34)... Anisodactylus Dejean 5(3) Male protarsi dilated laterally and spongily pubescent 6(3) Segment 4 of protarsi and mesotarsi with 2 ventrally (Fig. 12) ...(p. 54)... Pelmatellina (in part) membranous laminae (Fig. 25). Forebody (head and thorax) much narrower than elytra (Fig. 211–213, 221– — Male protarsi dilated laterally and biseriately pubescent 225) ...... 7 ventrally (Fig. 13) or unmodified ...... (p. 72)... Stenolophina — Segment 4 of protarsi and mesotarsi without membranous laminae (Fig. 26). Forebody (head and thorax) at most moderately narrower than elytra... 8 Alternative key to genera of New Zealand Harpalini 7(6) Elytral interneurs (Fig. 108) complete, consisting of Note. The key to subtribes provided above and keys to striae. Mentum with medial tooth as long as lateral genera within each subtribe allow the identification of all lobes (Fig. 16)...... (p. 68)... Syllectus Bates Harpalini genera, but because the key to subtribes is mainly — Elytral interneurs (Fig. 113) incomplete, consisting of based on males, an alternative key to genera, one bypass- rows of punctures. Mentum with medial tooth longer ing subtribes, is here provided for easier identification. than lateral lobes (Fig. 15) ...... 1 Rows of setiferous punctures present on elytral intervals ...... (p. 80)... Pholeodytes Britton 3, 5, or 7 (Fig. 97, 99), or on interneur 2 (Fig. 98) . 2 8(6) Eyes normally developed (Fig. 110). Mandibles shorter — Rows of setiferous punctures absent (Fig. 91) on elytral (Fig. 110) ...... 9 intervals 3, 5, and 7, and on interneur 2 ...... 3 — Eyes strongly reduced, flat or rather flat, consisting of 2(1) Metatarsomere 1 as long as metatarsomeres 2+3 (Fig. obliterated facets (Fig. 102, 112). Mandibles very long 188). Forebody (head and thorax) with sparse setiferous (about 5–6× their maximum width; Fig. 102, 112) 19 micropores dorsally (Fig. 136). Eyes strongly reduced, 9(8) Ventrites 5+6 with numerous short setae, in addition rather flat (Fig. 96–99). Tempora inflated (Fig. 96–99) to paired ambulatory setae (Fig. 28) ...... (p. 46)... Tuiharpalus new genus ...... (p. 73)... Euthenarus Bates — Metatarsomere 1 as long as metatarsomeres 2+3+4 (Fig. — Ventrites 5+6 without short setae, with paired 179). Forebody (head and thorax) without sparse ambulatory setae only (Fig. 27) ...... 10 26 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

10(9) Elytral interneurs incomplete basally and laterally — Body shape not boat-like, elytra not subtriangular (Fig. (Fig. 111). Pronotum suborbicular (Fig. 163) ...... 93-95). Scutellum visible (Fig. 93–95). Labrum strongly ...... (p. 77)... Haplanister Moore transverse, subrectangular, straight or slightly — Elytral interneurs complete (Fig. 104). Pronotum not emarginate apically (Fig. 93–95) ...... 18 suborbicular ...... 11 18(17) Body dark in colour. Tarsi pubescent dorsally. 11(10) Umbilicate setiferous series of elytral interval 9 Metafemora with 2 long setae on posterior margin. separated into 2 major groups (Fig. 107) ...... 12 Metatarsomere 1 as long as metatarsomeres 2+3 (Fig. 185) ...... (p. 42)... Parabaris Broun — Umbilicate setiferous series of elytral interval 9 not separated into 2 major groups (Fig. 93) ...... 17 — Body pale in colour. Tarsi glabrous dorsally. Metafemora with 5–7 long setae on posterior margin. Metatarsomere 12(11) Frons with clypeo-ocular prolongations (Fig. 107) 1 shorter than metatarsomeres 2+3 (Fig. 187) ...... 13 ...... (p. 45)... Triplosarus Bates — Frons without clypeo-ocular prolongations (Fig. 92) 19(8) Pronotum cordate or subcordate (Fig. 140–143)...... 14 Antennae widening from base to apex (Fig. 199); 13(12) Apex of prosternal lobe pubescent. Penultimate pubescence starting on antennomere 2 [NN–SD] ..... segment of labial palpi trisetose on anterior margin ...... (p. 54)... Hakaharpalus new genus (Fig. 10). Eyes widely separated from buccal fissure — Pronotum quadrate (Fig. 164). Antennae not widening ventrally (by 1.5–2.0× maximum width of antennal from base to apex (Fig. 220); pubescence starting on scape; Fig. 19). [North Island: ND] ...... antennomere 3 [TH] ...... (p. 57)... Kupeharpalus new genus ...... (p. 79)... Kiwiharpalus new genus — Apex of prosternal lobe glabrous (Fig. 2). Penultimate segment of labial palpi bisetose on anterior margin (Fig. 11). Eyes reaching buccal fissure (Fig. 21) or narrowly separated from it ventrally (by 0.3–0.7× maximum width of antennal scape; Fig. 20). [Throughout New Subtribe ANISODACTYLINA Zealand] ...... (p. 60)... Lecanomerus Chaudoir Diagnosis (New Zealand). Body length: 4.5–20.0 mm. 14(12) Metatarsomere 1 very long, almost as long as Frons without clypeo-ocular prolongations. Mentum usu- metatarsomeres 2+3+4 (Fig. 183) ...... ally with a tooth medially, seldom without a tooth ...... (p. 41)... Notiobia Perty (Anisodactylus, Gnathaphanus, Maoriharpalus). Mentum — Metatarsomere 1 much shorter, at most as long as, or and submentum usually separated by complete transverse slightly longer than, metatarsomeres 2+3 ...... 15 suture, seldom by laterally incomplete transverse suture 15(14) Elytra fused along suture; hindwings vestigial. (Gaioxenus), or without suture (Anisodactylus). Penulti- Pronotum not subrectangular (Fig. 114–120). mate segment of labial palpi usually plurisetose (with 4 Metafemora with 2 long setae on posterior margin ... setae or more) on anterior margin, seldom trisetose (with 3 ...... (p. 27)... Allocinopus Broun setae; Allocinopus angustulus, A. smithi, Hypharpax australis, Tuiharpalus clunieae, T. hallae). Apex of — Elytra free along suture; hindwings fully developed. prosternal lobe pubescent. Male protarsi dilated laterally Pronotum subrectangular (Fig. 124–125, 137–139). and spongily pubescent ventrally; male mesotarsi usually Metafemora with 4–10 long setae on posterior margin dilated laterally and spongily pubescent ventrally, seldom ...... 16 unmodified. Metatarsomere 1 of variable length. Umbili- 16(15) Metatarsomere 5 with 6–8 setae ventrally. Posterior cate setiferous series of interval 9 usually continuous, sel- bead of pronotum complete (Fig. 137–139). [Body dom separated into 2 major groups (Allocinopus, length 6–12 mm.] ...... (p. 51)... Harpalus Latreille Hypharpax, Notiobia) with posterior group continuous — Metatarsomere 5 with 4 setae ventrally. Posterior bead (not divided further into 2 subgroups). Aedeagus arcuate, of pronotum incomplete medially (Fig. 124–125). asymmetrical (with ostium deflected to the right, twisted [Body length 4.5–7.0 mm.] ...... or undulated) or symmetrical (with ostium dorsal, not de- ...... (p. 37)... Hypharpax Macleay flected laterally). 17(11) Body shape boat-like, with subtriangular elytra Geographic distribution. Worldwide. (Fig. 88). Scutellum hidden (Fig. 88). Labrum slightly References. Noonan 1973: 266–480 (classification; key transverse, almost square, convex apically (Fig. 88) . to genera) and 1976: 8–15 (taxonomy); Larochelle & ...... (p. 35)... Gaioxenus Broun Larivière 2001: 122–126 (catalogue). Fauna of New Zealand 53 27

Key to genera of New Zealand Anisodactylina — Body shape not boat-like, elytra not subtriangular (Fig. 1 Rows of setiferous punctures present on elytral intervals 93–95). Scutellum visible (Fig. 93–95). Labrum 3, 5, or 7 (Fig. 97, 99), or on interneur 2 (Fig. 98) . 2 strongly transverse, straight or slightly emarginate apically (Fig. 93–95) ...... 9 — Rows of setiferous punctures absent (Fig. 91) on elytral 9(8) Body pale in colour. Tarsi glabrous dorsally. intervals 3, 5, and 7, and on interneur 2 ...... 3 Metafemora with 5–7 long setae on posterior margin. 2(1) Metatarsomere 1 as long as metatarsomeres 2+3 (Fig. Paraglossae as long as ligula (Fig. 31) ...... 188). Forebody (head and thorax) with sparse setiferous ...... (p. 45)... Triplosarus Bates micropores dorsally (Fig. 136). Eyes strongly reduced, — Body dark in colour. Tarsi pubescent dorsally. rather flat (Fig. 96–99). Tempora inflated (Fig. 96–99) Metafemora with 2 long setae on posterior margin...... (p. 46)... Tuiharpalus new genus Paraglossae longer than ligula ...... — Metatarsomere 1 longer, as long as metatarsomeres ...... (p. 42)... Parabaris Broun 2+3+4 (Fig. 179). Forebody (head and thorax) without sparse setiferous micropores dorsally. Eyes moderately large, convex (Fig. 89). Tempora not inflated (Fig. 89) ...... (p. 36)... Gnathaphanus Macleay Genus Allocinopus Broun, 1903 E Allocinopus Broun, 1903: 607. Type species: Allocinopus 3(1) Mentum without a tooth medially (Fig. 18) ...... 4 sculpticollis Broun, 1903, by monotypy. — Mentum with a tooth medially (Fig. 14) ...... 5 Description. Body length: 6.0–11.5 mm. Forebody (head 4(3) Mandibles (Fig. 91) and antennal scapes (Fig. 182) and thorax) without sparse setiferous micropores dorsally. very long, about 6× their maximum width. Labrum Head. Mandibles moderately long, slightly curved for- strongly emarginate apically (Fig. 91). Mentum and ward, blunt apically. Labrum strongly transverse; apex submentum separated by transverse suture (Fig. 22). straight or slightly emarginate medially. Eyes moderately Pronotum suborbicular (Fig. 126). [TH] ...... large, slightly to strongly convex, widely separated from ...... (p. 40)... Maoriharpalus new genus buccal fissure ventrally (by about 2× maximum width of — Mandibles (Fig. 87) and antennal scapes (Fig. 177) antennal scape). Tempora not inflated. Frons without much shorter. Labrum straight or slightly emarginate clypeo-ocular prolongations. Antennal pubescence start- apically (Fig. 87). Mentum and submentum fused, not ing from basal third or half of antennomere 3. Mentum separated by transverse suture (Fig. 24). Pronotum with a tooth medially, moderately shorter than lateral lobes. rectangular (Fig. 121). [South Island and southern North Mentum and submentum separated by complete trans- Island]...... (p. 34)... Anisodactylus Dejean verse suture. Paraglossae as long as or longer than ligula. Palpi with last segment fusiform, seldom truncate apically 5(3) Umbilicate setiferous series of elytral interval 9 (latitarsis), with sparse, short or moderately long pubes- separated into 2 major groups (Fig. 85) ...... 6 cence; penultimate segment of labial palpi plurisetose or — Umbilicate setiferous series of elytral interval 9 not trisetose on anterior margin. Thorax. Pronotum cordate or separated into 2 major groups (Fig. 93) ...... 8 moderately transverse; base straight, as wide as or moder- 6(5) Elytra fused along suture; hindwings vestigial. ately narrower than elytral base; lateral beads complete; Pronotum moderately transverse (Fig. 114–120)...... anterior bead incomplete medially; posterior bead com- ...... (p. 27)... Allocinopus Broun plete or incomplete medially. Scutellum visible. Apex of — Elytra free along suture; hindwings fully developed. prosternal lobe pubescent. Legs. Metafemora with 2 long Pronotum very transverse (Fig. 124–125, 127) .... 7 setae on posterior margin. Male protarsi and mesotarsi dilated laterally and spongily pubescent ventrally. Seg- 7(6) Metatarsomere 1 parallel-sided, very long, almost as ment 4 of protarsi and mesotarsi of both sexes without long as metatarsomeres 2+3+4 (Fig. 183) ...... membranous laminae. Tarsi glabrous or pubescent (a few ...... (p. 41)... Notiobia Perty or numerous setae) dorsally; metatarsomere 5 pubescent — Metatarsomere 1 subtriangular, short, only about as (4-6 setae) ventrally; metatarsomere 1 as long as, shorter long as metatarsomere 2 (Fig. 181)...... or longer than metatarsomeres 2+3. Elytra. Interneurs com- ...... (p. 37)... Hypharpax Macleay plete. Rows of setiferous punctures absent on intervals 3, 8(5) Body shape boat-like, with subtriangular elytra (Fig. 5, and 7, and on interneur 2. Umbilicate setiferous series of 88). Scutellum hidden (Fig. 88). Labrum slightly interval 9 separated into two major groups, with posterior transverse, almost square, convex apically (Fig. 88) . group continuous. Abdomen. Ventrites 2+3 of male with- ...... (p. 35)... Gaioxenus Broun out a setiferous fovea. Ventrites 5+6 of both sexes without 28 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) short setae, with paired ambulatory setae only. Aedeagus. 5(4) Pronotum (Fig. 114): posterior angles subrectangular; Lateral view: slightly or strongly arcuate. Dorsal view: sides slightly sinuate. Aedeagus (Fig. 32, lateral view) symmetrical (with ostium dorsal, not deflected laterally) with apex straight; symmetrical (in dorsal view) [North or asymmetrical (with ostium deflected to the right); dor- Island, north of the Central Volcanic Plateau] ...... sal membranous area wide, extending almost to basal bulb; ...... (p. 28)... smithi Broun apical disc present. Internal sac armed. — Pronotum (Fig. 115): posterior angles obtuse; sides Geographic distribution. New Zealand (endemic). almost straight. Aedeagus (Fig. 33, lateral view) with References. Hudson, 1934: 177 (list); Noonan, 1973: 284– apex deflected dorsally; asymmetrical (in dorsal view) 285 and 1976: 8 (taxonomy); Larochelle & Larivière, 2001: [North Island, south of the Central Volcanic Plateau] 122–123 (catalogue)...... (p. 29)... angustulus Broun Remarks. Apart from variation in the pubescence of the 6(4) Base of pronotum (Fig. 117) much narrower than penultimate segment of the labial palpi, Allocinopus spe- elytral base; posterior angles obtuse; sides not sinuate. cies show a high degree of similarity in morphological char- Elytral shoulders rounded (Fig. 117). Aedeagus (Fig. acters, including the male genitalia, suggesting that they 35, dorsal view) with apical disc broadly spatulate .. form a distinct monophyletic group...... (p. 31)... bousqueti new species — Base of pronotum (Fig. 116) slightly narrower than elytral base; posterior angles subrectangular; sides Key to species of Allocinopus slightly sinuate. Elytral shoulders angulate (Fig. 116). Aedeagus (Fig. 34, dorsal view) with apical disc 1 Metatarsomere 1 short, only slightly longer than narrowly spatulate ...... (p. 30)... belli new species metatarsomere 2 (Fig. 175). Palpi truncate apically. Male protarsi and mesotarsi strongly dilated laterally (each tarsomere about 2× wider than long; Fig. 175). E [Pronotum (Fig. 119). Chatham Islands] ...... Allocinopus smithi Broun, 1912 ...... (p. 32)... latitarsis Broun Figures 32, 85, 114, 170; Map p. 147 Allocinopus smithi Broun, 1912: 391. Lectotype (here desig- — Metatarsomere 1 as long as (Fig. 170) or longer (Fig. nated): female (BMNH) labelled “paratype (circular red- 173) than metatarsomeres 2+3. Palpi not truncate bordered label; typed) / 3177. (hand-written) / New Zea- apically. Male protarsi and mesotarsi not strongly land. Broun Coll. Brit. Mus. 1922-482. (typed) / dilated laterally (Fig. 176) ...... 2 Ratapihipihi. Nov. 1909. (hand-written) / Allocinopus 2(1) Pronotum cordate (heart-shaped), sides clearly sinuate smithi. female. (hand-written) / LECTOTYPE Allocinopus smithi Broun, 1912 designated by Larochelle (Fig. 118, 120). Metepisterna longer than wide ..... 3 & Larivière 2004 (red label; typed).” Fair condition. — Pronotum not cordate, sides little or not sinuate (Fig. Posterior legs without metatarsi. Paralectotype: 1 male 114–117). Metepisterna as wide as or wider than long (BMNH) bearing blue paralectotype label...... 4 Allocinopus castaneus Broun, 1912: 392. Lectotype (here designated): female (BMNH) labelled “Type (circular red- 3(2) Pronotum (Fig. 118): punctuation weakly developed bordered label; typed) / 3178. female (hand-written) / basally and laterally; basal foveae deep, much longer Hunua. Maketu. (hand-written) / New Zealand. Broun than wide. Body brownish. Tarsi pubescent dorsally Coll. Brit. Mus. 1922-482. (typed) / Allocinopus ...... (p. 31)... wardi new species castaneus. female. (hand-written) / LECTOTYPE Allocinopus castaneus Broun, 1912 designated by — Pronotum (Fig. 120): punctuation strongly developed Larochelle & Larivière 2004 (red label; typed).” Perfect basally and laterally; basal foveae shallow, about as condition. New synonym. long as wide. Body blackish. Tarsi glabrous dorsally Description. Body length: 6.5–7.0 mm. Moderately con- ...... (p. 33)... sculpticollis Broun vex. Reddish black; head darker; lateral margins of pronotum 4(2) Penultimate segment of labial palpi trisetose (with 3 and elytra paler; antennae, palpi, and legs light reddish. setae) on anterior margin (Fig 10). Metepisterna wider Generally glabrous and smooth. Microsculpture than long. Metatarsomere 1 as long as metatarsomeres isodiametric and strong on head, moderately transverse 2+3 (Fig. 170) ...... 5 and weak on pronotum, very transverse (with microlines) — Penultimate segment of labial palpi plurisetose (with 4 and weak on elytra (in males) or very transverse (with setae or more) on anterior margin (Fig. 9). Metepisterna microlines) and strong (in females). Shiny, without metal- as wide as long. Metatarsomere 1 longer than lic lustre. Head. Moderately large, narrower across eyes metatarsomeres 2+3 (Fig. 173) ...... 6 than pronotal apex; flat anteriorly, slightly convex Fauna of New Zealand 53 29 posteriorly. Labrum with apex slightly emarginate medi- 2001: 123 (including castaneus; taxonomy, distribution, ally. Eyes moderately large, slightly convex. Antennae ecology, biology, dispersal power). moderately long, reaching pronotal base; antennal scape Remarks. Broun described smithi from a pair of “muti- about 2× longer than its maximum width; pubescence start- lated specimens”, one of which (a female) is here desig- ing from basal 1/3 of antennomere 3. Paraglossae as long as nated as lectotype. The second specimen represented in ligula. Palpi not truncate apically, with sparse, moderately the Natural History Museum, London (BMNH) collec- long pubescence; penultimate segment of labial palpi tion is a perfectly preserved male specimen with label trisetose (with 2 long setae and 1 short seta) on anterior information fitting the original description; therefore, it is margin. Thorax. Pronotum (Fig. 114) very transverse, considered to be the paralectotype. Broun described widest before middle; sides slightly converging toward base, castaneus from two females, one of which could be located slightly sinuate; base straight, as wide as elytral base; apex in the Natural History Museum, London (BMNH) and is concave; lateral depressions widening posteriorly; poste- here designated as lectotype. These two lectotypes are rior bead complete medially; anterior angles strongly de- designated to preserve stability of nomenclature in the veloped, acute; posterior angles strongly developed, future. The lectotype of castaneus although pale in colour subrectangular; basal foveae shallow, narrow; anterior lat- conforms morphologically with smithi; the two taxa are eral setiferous puncture not touching lateral beads; punc- believed to be conspecific. Allocinopus smithi resembles tuation feebly developed. Apex of prosternal lobe with 1- angustulus in its general morphology and its trisetose palpi 2 long setae and several short setae. Metepisterna wider (plurisetose in other Allocinopus species). The two spe- than long. Legs. Tarsi pubescent (with numerous setae) cies are highly variable morphologically (within and be- dorsally; metatarsomere 5 pubescent (with 6 setae) tween populations) and the only reliable diagnostic char- ventrally; metatarsomere 1 as long as metatarsomeres 2+3. acter between them is the male aedeagus. The geographic Elytra. Widest about middle. Shoulders strongly devel- distribution of these two allopatric species may also assist oped, angulate, with a tooth. Subapical sinuations feeble. their identification. Sutural apices angulate-rounded. Scutellar striole present. Interneurs shallow, deepening apically, impunctate. Inter- vals sparsely punctate, flat, becoming convex apically. In- Allocinopus angustulus Broun, 1912 E terval 3 without setiferous puncture behind middle. Figures 33, 115, 171; Map p. 147 Aedeagus (Fig. 32). Lateral view: strongly arcuate; apex Allocinopus angustulus Broun, 1912: 392. Holotype: female narrowly pointed, straight. Dorsal view: symmetrical (with (BMNH) labelled “Type (circular red-bordered label; ostium dorsal, not deflected laterally); dorsal membranous typed) / 3179. female. (hand-written) / New Zeal. Broun area very wide, extending almost to basal bulb; apex straight Coll. Brit. Mus. 1922-482 (typed) / Forty Mile-Bush. (not deflected laterally) and apical disc present, truncate at Suter.. (hand-written) / Allocinopus angustulus.” Perfect tip, with divergent sides. Internal sac armed. condition. Material examined. 421 specimens, including types Description. Body length: 6.0–7.5 mm. Moderately con- (AMNZ, BMNH, ITNZ, JNNZ, LUNZ, MONZ, NZAC, vex. Reddish black; head darker; lateral margins of pronotum UCNZ). and elytra, sutural intervals rufous; antennae, palpi, and legs light reddish. Generally glabrous and smooth. Geographic distribution (Map p. 147). North Island: Microsculpture isodiametric and strong on head, moder- AK, BP, TK, TO, WO. ately transverse and weak on pronotum, very transverse Ecology. Lowland, montane. Wet forests (broadleaf, (with microlines) on elytra. Shiny, without metallic lustre. podocarp): along streams, gullies, and mud flats. Shaded Head. Moderately large, narrower across eyes than pronotal ground; soil covered with dead leaves. Nocturnal; shelter- apex; flat anteriorly, slightly convex posteriorly. Labrum ing during the day under stones, in leaf litter, under fallen with apex slightly emarginate medially. Eyes moderately branches and logs. Gregarious. Biology. Seasonality: Sep- large, slightly convex. Antennae moderately long, reaching tember–January, March–April, June, August. Tenerals: pronotal base; antennal scape about 2× longer than its maxi- September–October, December, March–April. Occasion- mum width; pubescence starting from basal third of ally infested with fungi (Laboulbeniales). Dispersal power. antennomere 3. Paraglossae as long as ligula. Palpi not Elytra fused along suture. Subapterous. Moderate runner. truncate apically, with sparse, moderately long pubescence; Good burrower. Collecting techniques. Turning stones penultimate segment of labial palpi trisetose (with 2 long and pieces of wood; raking leaf litter; pitfall trapping. setae and 1 short seta) on anterior margin. Thorax. References. Hudson, 1934: 37 (distribution, ecology); Pronotum (Fig. 115) strongly transverse, widest before Noonan, 1973: 285 (taxonomy); Larochelle & Larivière, middle; sides slightly converging toward base, not sinuate, 30 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) almost straight; base straight, as wide as elytral base; apex Description. Body length: 7.5–8.0 mm. Moderately con- concave; lateral depressions widening posteriorly; poste- vex. Dark brownish; head darker; antennae and legs (except rior bead complete medially; anterior angles strongly de- femora) dark reddish; apical half of tibia light yellowish veloped, acute; posterior angles strongly developed, ob- brown; sides of pronotum and apical half of elytra reddish tuse; basal foveae deep, narrow; anterior lateral setiferous brown. Generally glabrous and smooth. Microsculpture punctures not touching lateral beads; punctuation feebly moderately strong, isodiametric on head, moderately trans- developed. Apex of prosternal lobe with 1–2 long setae verse on pronotum, and strongly transverse (with and several short setae. Metepisterna wider than long. Legs. microlines) on elytra. Head slightly shiny, pronotum mod- Tarsi pubescent (with numerous setae) dorsally; erately shiny, elytra very shiny and iridescent; without metatarsomere 5 pubescent (with 6 setae) ventrally; metallic lustre. Head. Moderately large, narrower across metatarsomere 1 as long as metatarsomeres 2+3. Elytra. eyes than pronotal apex; flat anteriorly, slightly convex Widest about middle. Shoulders strongly developed, posteriorly. Labrum with apex straight or slightly emarginate angulate, with a tooth. Subapical sinuations feeble. Sutural medially. Eyes moderately large, slightly convex. Anten- apices angulate. Scutellar striole present. Interneurs shal- nae moderately long, reaching elytral base; antennal scape low, deepening apically, impunctate. Intervals sparsely about 2× longer than its maximum width; pubescence start- punctate, flat, becoming convex apically. Interval 3 with- ing from middle of antennomere 3. Paraglossae longer than out setiferous puncture behind middle. Aedeagus (Fig. ligula. Palpi not truncate apically, with sparse, moderately 33). Lateral view: strongly arcuate; apex narrowly pointed, long pubescence; penultimate segment of labial palpi slightly deflected dorsally. Dorsal view: asymmetrical (with plurisetose (with 2–3 long setae and 4 short setae) on ostium slightly deflected to the right); dorsal membranous anterior margin. Thorax. Pronotum (Fig. 116) strongly area very wide, extending almost to basal bulb; apex de- transverse, widest before middle; sides slightly converging flected to the right and apical disc present, rounded at tip, toward base, slightly sinuate; base straight, slightly nar- almost parallel-sided. Internal sac armed. rower than elytral base; apex concave; lateral depressions Material examined. 143 specimens, including type widening posteriorly; posterior bead complete medially; (BMNH, ITNZ, JNNZ, MONZ, NZAC, UCNZ). anterior angles strongly developed, acute; posterior angles strongly developed, subrectangular; basal foveae deep, nar- Geographic distribution (Map p. 147). North Island: row; anterior lateral setiferous punctures not touching lat- HB, RI, WA, WI, WN. eral beads; punctuation feebly developed. Apex of Ecology. Lowland, montane. Wet or moist forests prosternal lobe with 2 long setae and several short setae. (broadleaf, beech): along streams, rills, and gullies. Shaded Metepisterna as wide as long. Legs. Tarsi pubescent (with ground; soil covered with dead leaves. Nocturnal; shelter- numerous setae) dorsally; metatarsomere 5 pubescent (with ing during the day under stones (mostly) and in leaf litter. 4 setae) ventrally; metatarsomere 1 longer than Gregarious. Biology. Seasonality: October–March, June– metatarsomeres 2+3. Elytra. Widest about middle. Shoul- August. Tenerals: February–March. Dispersal power. ders strongly developed, angulate, without a tooth. Sub- Elytra fused along suture. Subapterous. Moderate runner. apical sinuations feeble. Sutural apices angulate. Scutellar Good burrower. Collecting techniques. Turning stones, striole absent or present. Interneurs shallow, not deepen- pitfall trapping. ing apically, impunctate. Intervals impunctate, flat, not References. Noonan, 1973: 285 (taxonomy); Larochelle convex apically. Interval 3 without setiferous puncture & Larivière, 2001: 122 (taxonomy, distribution, ecology, behind middle. Aedeagus (Fig. 34). Lateral view: strongly biology, dispersal power). arcuate; apex narrowly rounded, slightly deflected dorsally. Remarks. See under A. smithi. Dorsal view: asymmetrical (with ostium slightly deflected to the right); dorsal membranous area moderately wide, extending almost to basal bulb; apex deflected to the right Allocinopus belli new species E and apical disc present, narrowly spatulate. Internal sac Figures 34, 116, 172; Map p. 147 armed. Allocinopus belli Larochelle & Larivière, new species. Material examined. 13 specimens, including types Holotype: male (NZAC) labelled “Whanarua Bay, Bay (BBNZ, ITNZ, LUNZ, NZAC). of Plenty under stone 24-i-62 J.C. Watt (hand-written) / J.C. Watt Collection Ent. Div. DSIR, 1966. (typed) / Geographic distribution (Map p. 147). North Island: HOLOTYPE [male symbol] Allocinopus belli new spe- BP–Papatea. Whanarua Bay. cies Larochelle & Larivière, 2004 (red label; typed).” Ecology. Lowland (coastal). Wet forests (broadleaf): along Paratypes: 1 female (NZAC), 4 males (MONZ; mounted on same card) from the same locality as the holotype, streams. Shaded ground; soil covered with dead leaves. bearing blue paratype labels. Nocturnal; sheltering during the day under stones. Biol- Fauna of New Zealand 53 31 ogy. Seasonality: October–November, January. Tenerals: numerous setae) dorsally; metatarsomere 5 pubescent (with late January. Dispersal power. Elytra fused along suture. 4 setae) ventrally; metatarsomere 1 longer than Subapterous. Moderate runner. Good burrower. Collect- metatarsomeres 2+3. Elytra. Widest about middle. Shoul- ing techniques. Turning stones. ders strongly developed, rounded, without a tooth. Sub- Remarks. This species closely resembles A. bousqueti but apical sinuations absent. Sutural apices rounded. Scutellar can be most easily distinguished from it by characters of striole absent or present. Interneurs shallow, deepening the male genitalia. In addition, A. belli is typically a coastal apically, impunctate. Intervals impunctate, flat, not con- species whereas A. bousqueti is an inland species. This vex apically. Interval 3 without setiferous puncture behind taxon is named after our friend and colleague Ross T. Bell middle. Aedeagus (Fig. 35). Lateral view: strongly arcu- (University of Vermont, Burlington) for his special help ate; apex spatulate, slightly deflected dorsally. Dorsal view: and encouragement in our carabid studies. asymmetrical (with ostium moderately deflected to the right); dorsal membranous area moderately wide, extend- ing almost to basal bulb; apex deflected to the right and Allocinopus bousqueti new species E broadly spatulate; apical disc present. Internal sac armed. Figures 35, 117, 173; Map p. 147 Material examined. 16 specimens, including types Allocinopus bousqueti Larochelle & Larivière, new species. (JNNZ, LUNZ, NZAC). Holotype: male (NZAC) labelled “NEW ZEALAND BP Waioeka Gorge Mangapumarumaru Track 100m Geographic distribution (Map p. 147). North Island: 25.XI.1997 Larochelle, Larivière (typed) / Moist BP–Ruatoria State Forest. Urewera National Park, Waimana broadleaf forest: under stones. (typed) / HOLOTYPE Valley, Te Waiiti Stream. Waioeka Gorge. Whinray Scenic [male symbol] Allocinopus bousqueti Larochelle & Reserve. GB–Morere Springs Scenic Reserve. Larivière, 2004 (red label; typed).” Paratypes: 2 males (1 NZAC, 1 MONZ) from the same locality as the Ecology. Lowland. Wet or moist forests (broadleaf): along holotype, bearing blue paratype labels. streams and mud flats. Shaded ground; soil covered with Description. Body length: 6.5–8.0 mm. Moderately con- dead leaves. Nocturnal; sheltering during the day under vex. Dark brownish; head darker; antennae and tarsi dark logs, fallen branches and epiphyte crowns, and in leaf lit- reddish; femora brownish; mandibles mostly brownish; ter. Biology. Seasonality: October–November, January, sides of pronotum reddish brown. Generally glabrous and April. Tenerals: November, April. Dispersal power. Elytra smooth. Microsculpture moderately strong, isodiametric fused along suture. Subapterous. Moderate runner. Good on head, moderately transverse on thorax, and strongly burrower. Collecting techniques. Turning stones and transverse (with microlines) on elytra. Head slightly shiny, pieces of wood. pronotum moderately shiny, elytra very shiny and irides- Remarks. This taxon is named after our close friend and cent; without metallic lustre. Head. Moderately large, nar- colleague Yves Bousquet (Agriculture and Agri-Food rower across eyes than pronotal apex; flat anteriorly, slightly Canada, Ottawa, Canada) for his special help and encour- convex posteriorly. Labrum with apex straight or slightly agement in our carabid studies. See also Remarks under A. emarginate medially. Eyes moderately large, slightly con- belli. vex. Antennae moderately long, reaching elytral base; antennal scape about 2× longer than its maximum width; Allocinopus wardi new species E pubescence starting from middle of antennomere 3. Figures 36, 118, 174; Map p. 147 Paraglossae longer than ligula. Palpi not truncate apically, Allocinopus wardi Larochelle & Larivière, new species. with sparse, moderately long pubescence; penultimate seg- Holotype: male (NZAC) labelled “NEW ZEALAND CL ment of labial palpi plurisetose (with 2–3 long setae and 4 Moehau Ra, Fantail Ck 24.XII. 1993 Larochelle, Larivière short setae) on anterior margin. Thorax. Pronotum (Fig. (typed) / Stony-gravelly stm edge, forested gully 117) very transverse, widest before middle; sides slightly (kohekohe/pepper tree). Under stones. (typed) / converging toward base, not sinuate; base straight, moder- HOLOTYPE [male symbol] Allocinopus wardi Larochelle ately narrower than elytral base; apex concave; lateral de- & Larivière, 2004 (red label; typed).” pressions widening posteriorly; posterior bead complete Description. Body length: 5.0 mm (only one specimen medially; anterior angles strongly developed, acute; poste- seen, head missing). Slightly convex. Brownish; sides of rior angles moderately developed, obtuse; basal foveae shal- pronotum and elytra reddish brown. Generally glabrous low, narrow; anterior lateral setiferous punctures touching and smooth. Microsculpture strong, very transverse (with lateral beads; punctuation feebly developed. Apex of microlines) on pronotum and elytra. Very shiny, without prosternal lobe with 2 long setae and several short setae. metallic lustre. Iridescent. Head. [Missing]. Thorax. Metepisterna as wide as long. Legs. Tarsi pubescent (with Pronotum (Fig. 118) cordate (heart-shaped), widest before 32 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) middle; sides strongly converging toward base, strongly Description. Body length: 7.5–11.5 mm. Slightly convex. sinuate; base straight, slightly narrower than elytral base; Brownish; head darker; antennae and tarsi reddish or yel- apex concave; lateral depressions widening posteriorly; lowish; apical half of tibiae light yellowish brown; mandi- posterior bead incomplete medially; anterior angles strongly bles mostly brownish; sides of pronotum and elytra red- developed, acute; posterior angles strongly developed, dish brown. Generally glabrous and smooth. Microsculpture subrectangular; basal foveae deep, narrow, very long; ante- absent or barely visible on head and pronotum, weak and rior lateral setiferous punctures touching lateral beads; moderately transverse on male elytra, strong and punctuation feebly developed. Apex of prosternal lobe isodiametric on female elytra. Very shiny, without metallic with 2 long setae and several short setae. Metepisterna lustre. Head. Moderately large, as wide across eyes as longer than wide. Legs. Tarsi pubescent (with several se- pronotal apex; flat anteriorly, slightly convex posteriorly. tae) dorsally; metatarsomere 5 pubescent (with 4 setae) Labrum with apex straight or slightly emarginate medially. ventrally; metatarsomere 1 as long as metatarsomeres 2+3. Eyes very large and convex. Antennae moderately long, Elytra. Widest about middle. Shoulders strongly devel- reaching elytral base; antennal scape about 2× longer than oped, angulate, without a tooth. Subapical sinuations ab- its maximum width; pubescence starting from middle of sent. Sutural apices angulate. Scutellar striole present. antennomere 3. Paraglossae longer than ligula. Palpi trun- Interneurs shallow, impunctate. Intervals impunctate, flat, cate apically, with sparse short pubescence; penultimate becoming convex apically. Interval 3 without setiferous segment of labial palpi plurisetose (with 3 long setae and 4 puncture behind middle. Aedeagus (Fig. 36). Lateral view: short setae) on anterior margin. Thorax. Pronotum (Fig. strongly arcuate; apex blunt, sinuate, deflected ventrally. 119) very transverse, widest before middle; sides strongly Dorsal view: asymmetrical (with ostium slightly deflected converging toward base, not sinuate; base straight, moder- to the right); dorsal membranous area moderately wide, ately narrower than elytral base; apex concave; lateral de- extending almost to basal bulb; apex deflected to the right; pressions widening posteriorly; posterior bead complete; apical disc present, rounded-triangular. Internal sac armed. anterior angles moderately developed, obtuse; posterior Material examined. Holotype (NZAC). angles moderately developed, rounded; basal foveae deep, wide; anterior lateral setiferous punctures touching lateral Geographic distribution (Map p. 147). North Island: beads; punctuation feebly developed. Apex of prosternal CL–Moehau Range, Fantail Creek. lobe with 2 long setae and several short setae. Metepisterna Ecology. Lowland. A wet broadleaf forest: along a stream. longer than wide. Legs. Tarsi pubescent (with a few setae) Shaded ground; soil stony-gravelly, covered with dead dorsally; metatarsomere 5 pubescent (with 4 setae) leaves. Found dead under a stone. Biology. Seasonality: ventrally; metatarsomere 1 short, slightly longer than unknown. Dispersal power. Elytra fused along suture. metatarsomere 2. Male protarsi and mesotarsi strongly Subapterous. Moderate runner (after leg morphology). dilated laterally, about 2× wider than long (contrary to Remarks. The configuration of the male aedeagus is unique other Allocinopus species). Elytra. Widest about middle. among Allocinopus species. This is the only species so far Shoulders strongly developed, rounded, without a tooth. recorded from the Coromandels and it seems geographi- Subapical sinuations feeble. Sutural apices angulate. cally restricted to that region. This species is named after Scutellar striole present. Interneurs shallow, not deepening our good friend and colleague John Ward (Canterbury apically, impunctate. Intervals impunctate, flat, not con- Museum, Christchurch) for his special encouragement to- vex apically. Interval 3 without setiferous puncture behind ward the establishment of our new life and career in New middle. Aedeagus (Fig. 37). Lateral view: slightly arcuate; Zealand. apex narrowly triangular, almost straight. Dorsal view: asymmetrical (with ostium slightly deflected to the right); dorsal membranous area very wide, extending almost to E Allocinopus latitarsis Broun, 1911 basal bulb; apex slightly deflected to the right; apical disc Figures 37, 119, 175; Map p. 147 present, triangular. Internal sac armed. Allocinopus latitarsis Broun, 1911: 95. Lectotype (here des- ignated): male (BMNH) labelled “51.male. (rectangular Material examined. 274 specimens, including types label; hand-written) / Chathams Is. Broun Coll. B.M. 1922- (AMNZ, BBNZ, BMNH, CMNZ, ITNZ, LUNZ, 482. (typed) / Pitt Island. -T. Hall- (hand-written) / MONZ, NZAC, UCNZ). Allocinopus latitarsis male. (hand-written) / Geographic distribution (Map p. 147). Offshore Islands: LECTOTYPE Allocinopus latitarsis Broun, 1911 desig- CH–several localities on Chatham Island, Pitt Island, South nated by Larochelle & Larivière 2004 (red label; typed). “ Good condition. Right middle leg without last tarsomere. East Island, North East Island, The Sisters. Paralectotypes: 2 males and 1 female (BMNH) bearing Ecology. Lowland. Wet forests (broadleaf), shrublands, blue paralectotype labels. and scrublands. Also pastures, gardens, stream edges, and Fauna of New Zealand 53 33 coastal rocky faces. Mostly shaded ground; soil usually ally. Eyes moderately large and convex. Antennae moder- covered with dead leaves. Nocturnal; sheltering during the ately long, reaching elytral base; antennal scape about 2× day under logs and stones (mostly), under fallen bark, longer than its maximum width; pubescence starting from plant debris, in leaf litter; in and around bird nests middle of antennomere 3. Paraglossae longer than ligula. (Puffinus). Gregarious. Biology. Seasonality: September– Palpi not truncate apically, with sparse short pubescence; March, July-August. Mating: September. Tenerals: Febru- penultimate segment of labial palpi plurisetose (with 3 ary. Occasionally infested with mites and fungi long setae and 4 short setae) on anterior margin. Thorax. (Laboulbeniales). Dispersal power. Elytra fused along Pronotum (Fig. 120) cordate (heart-shaped), widest before suture. Subapterous. Moderate runner. Good burrower. middle; sides strongly converging toward base, strongly Occasional climber (on tree trunks at night). Collecting sinuate; base straight, slightly narrower than elytral base; techniques. Turning logs and stones, pitfall trapping. apex concave; lateral depressions widening posteriorly References. Noonan, 1973: 285 (taxonomy); Watt, 1980: (more than in other Allocinopus species); posterior bead 334 (distribution, ecology); Emberson, 1998: 30; Larochelle complete; anterior angles strongly developed, acutely & Larivière, 2001: 122 (taxonomy, distribution, ecology, rounded; posterior angles strongly developed, biology, dispersal power). subrectangular; basal foveae shallow, about as long as wide; anterior lateral setiferous punctures touching lateral beads; Remarks. The lectotype of A. latitarsis is designated to punctuation strongly developed (basally and laterally). preserve stability of nomenclature in the future. The Apex of prosternal lobe with 2 long setae and several short aedeagus of this species is morphologically close to that of setae. Metepisterna longer than wide. Legs. Tarsi glabrous A. sculpticollis, but A. latitarsis is unique among Allocinopus dorsally; metatarsomere 5 pubescent (with 4 setae) species in having male protarsi and mesotarsi strongly di- ventrally; metatarsomere 1 as long as metatarsomeres 2+3. lated laterally, and metatarsomeres 1 very short. It is also Elytra. Widest about middle. Shoulders strongly devel- the only Allocinopus species so far recorded from the oped, angulate, without a tooth. Subapical sinuations fee- Chatham Islands and endemic to those islands. The exter- ble. Sutural apices angulate. Scutellar striole present. nal morphology of this species is highly variable, e.g., body Interneurs shallow, deepening apically, impunctate. Inter- shape and size can vary even within populations. vals impunctate, flat, becoming convex apically. Interval 3 without setiferous puncture behind middle. Aedeagus (Fig. Allocinopus sculpticollis Broun, 1903 E 38). Lateral view: slightly arcuate; apex narrowly pointed, Figures 38, 86, 120, 176; Map p. 147 slightly deflected ventrally. Dorsal view: symmetrical (with Allocinopus sculpticollis Broun, 1903: 608. Holotype: male ostium dorsal, not deflected laterally); dorsal membranous (BMNH) labelled “Type (circular red-bordered label; area very wide, extending almost to basal bulb; apex straight typed) / 2624 (hand-written) / male (hand-written) / (not deflected laterally); apical disc present, rounded-tri- New Zeal. Broun Coll. Brit. Mus. 1922-482 (typed) / angular. Internal sac armed. Motueka. Nelson. (hand-written) / Allocinopus Material examined. 422 specimens, including types sculpticollis. (hand-written)”. Good condition. Left pos- terior leg without last tarsomere. (AMNZ, BMNH, ITNZ, JNNZ, LUNZ, MONZ, NZAC, Allocinopus ocularius Broun, 1908: 344. Holotype: male UCNZ). (BMNH): labelled “Type (circular red-bordered label; Geographic distribution (Map p. 147). North Island: typed) / 2625 (hand-written) / New Zeal. Broun Coll. BP, GB, HB, RI, TK, TO, WA, WI, WN. South Island: Brit. Mus. 1922-482 (typed) / Manawatu near Gorge. (hand-written) / Allocinopus ocularius. male. (hand-writ- BR, NN, SD, WD. ten)”. Good condition. Left middle and left posterior legs Ecology. Lowland, montane. Wet forests (broadleaf, missing. New synonym. podocarp, beech), swamp forests, and shrublands: along Description. Body length: 9.0–11.0 mm. Moderately con- streams and mud flats. Shaded ground; soil covered with vex. Blackish (contrary to other Allocinopus species); an- dead leaves. Nocturnal; sheltering during the day in bur- tennae, palpi, and legs brownish red; mandibles mostly rows dug under stones (mostly) and logs, also hiding under dark red; head with 1–2 reddish spots on disc (contrary to fallen branches and epiphyte crowns, and in leaf litter. other Allocinopus species). Generally glabrous and smooth. Gregarious. Biology. Seasonality: October–April. Tenerals: Microsculpture strong, transverse (North Island) or October–April. Occasionally infested with mites and fungi isodiametric (South Island). Moderately shiny, without (Laboulbeniales). Dispersal power. Elytra fused along metallic lustre. Head. Moderately large, as wide across suture. Subapterous. Moderate runner. Good burrower. eyes as pronotal apex; flat anteriorly, slightly convex Collecting techniques. Turning stones and logs, pitfall posteriorly. Labrum with apex slightly emarginate medi- trapping. 34 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

References. Hudson, 1934: 37 (distribution, ecology); branous area wide, not extending to basal bulb; apical disc Noonan, 1973: 285 (taxonomy); Townsend, 1997: 16 and present. Internal sac unarmed. 1998: 11, 21 (distribution); Larochelle & Larivière, 2001: Geographic distribution. North America, Europe, Asia, 122–123 (including ocularius; taxonomy, distribution, ecol- northern Africa; New Zealand (adventive). ogy, biology, dispersal power). References. Noonan, 1973: 349–380 and 1976: 13 (tax- Remarks. Examination of the holotype of A. ocularius onomy); Larochelle & Larivière, 2001: 123 (catalogue). revealed it to be conspecific with A. sculpticollis. Body shape and size are highly variable within and between populations, throughout the range of this species. North Subgenus Anisodactylus Dejean, 1829 A Island and South Island populations show different types of body microsculpture, but aedeagal characters are con- Geographic distribution. Same as genus. stant across all populations examined. See also Remarks under A. latitarsis. Anisodactylus binotatus (Fabricius, 1787) A Figures 39, 87, 121, 177; Map p. 147 Genus Anisodactylus Dejean, 1829 A Carabus 2notatus Fabricius, 1787: 199. Type locality: Kiel, Germany. Anisodactylus Dejean 1829: 4. Type species: Carabus Multiple synonyms exist in the Old World literature for this binotatus Fabricius, 1787, designated by Westwood, 1838: adventive species. 4. Description (Anisodactylus binotatus). Body length: 10.0– Description. Body length: 10.0–12.7 mm. Moderately 12.7 mm. Forebody (head and thorax) without sparse convex. Black; antennae (segments 1–2), palpi (in part), setiferous micropores dorsally. Head. Mandibles moder- and legs rufous; frons with 2 rufous, more or less confluent ately long, slightly curved forward, blunt apically. Labrum spots medially. Generally glabrous and smooth; elytra with outermost intervals and apex of other intervals pubescent strongly transverse; apex straight or slightly emarginate medially. Eyes moderately large, convex, moderately sepa- and punctate. Microsculpture isodiametric (head, elytra), rated from buccal fissure ventrally (by about maximum with somewhat transverse meshes (pronotum). Shiny, with- width of antennal scape). Tempora not inflated. Frons with- out metallic lustre. Head. Moderately large, narrower across out clypeo-ocular prolongations. Antennal pubescence eyes than pronotal apex; flat anteriorly, slightly convex posteriorly. Antennae moderately long, reaching about starting from basal third of antennomere 3. Mentum with- × out a tooth medially. Mentum and submentum fused, not pronotum base; antennal scape about 2 longer than its maximum width. Penultimate segment of labial palpi with separated by transverse suture. Paraglossae longer than 6–7 long setae on anterior margin. Thorax. Pronotum (Fig. ligula. Palpi with last segment fusiform, not truncate 121) very transverse, widest about middle; sides converg- apically, with sparse, short pubescence; penultimate seg- ing toward base, not sinuate; apex concave; lateral depres- ment of labial palpi plurisetose on anterior margin. Tho- rax. Pronotum transverse, rectangular; base straight, as sions widening posteriorly; anterior angles strongly devel- wide as elytral base; lateral beads complete; anterior and oped, rounded; posterior angles strongly developed, posterior beads complete. Scutellum visible. Apex of subrectangular, with a tooth; basal foveae deep, wide; punc- prosternal lobe pubescent. Legs. Metafemora with 3–4 tuation strongly developed (basally and laterally). Apex of long setae on posterior margin. Male protarsi and mesotarsi prosternal lobe with 6–10 long setae and 3–6 short setae. dilated laterally and spongily pubescent ventrally. Seg- Metepisterna longer than wide. Elytra. Widest about mid- ment 4 of protarsi and mesotarsi of both sexes without dle. Shoulders strongly developed, rounded, without a membranous laminae. Tarsi glabrous (except metatarsomeres tooth. Subapical sinuations feeble. Sutural apices rounded. 1+2 with 3–6 setae) dorsally; metatarsomere 5 pubescent Scutellar striole present. Interneurs shallow, impunctate. (with 8 setae) ventrally; metatarsomere 1 as long as Intervals flat; outer intervals and apex of elytra finely and metatarsomeres 2+3. Elytra. Interneurs complete. Rows densely punctate. Interval 3 with setiferous puncture be- of setiferous punctures absent on intervals 3, 5, and 7, and hind middle. Aedeagus (Fig. 39). Lateral view: slightly on interneur 2. Umbilicate setiferous series of interval 9 arcuate, very long. Dorsal view: asymmetrical (with os- continuous. Abdomen. Ventrites 2+3 of male without a tium slightly deflected to the left), twisted at middle; dor- setiferous fovea. Ventrites 5+6 of both sexes without short sal membranous area wide, divided into two small parts; setae, with paired ambulatory setae only. Aedeagus. Lat- apical disc present. Internal sac unarmed. eral view: slightly arcuate. Dorsal view: asymmetrical (with Material examined. 31 non-type specimens (AMNZ, ostium deflected to the left), twisted at middle; dorsal mem- ITNZ, JNNZ, LUNZ, MONZ, NZAC, OMNZ). Fauna of New Zealand 53 35

Geographic distribution (Map p. 147). North Island: den (contrary to other Harpalini genera). Apex of prosternal WN. South Island: CO, DN, MC, OL, SC. Introduced lobe pubescent. Legs. Metafemora with 2 long setae on from Europe. First New Zealand record: Spreydon, posterior margin. Male protarsi and mesotarsi dilated lat- Christchurch, MC, 1938 (Pilgrim, 1963: 837). Well estab- erally and spongily pubescent ventrally. Segment 4 of lished. protarsi and mesotarsi of both sexes without membranous Ecology. Lowland, upland. Vicinities of cities. Gardens, laminae. Tarsi pubescent (with numerous setae) dorsally; cultivated fields (maize), pastures, tussock grasslands, dried metatarsomere 5 pubescent (with numerous setae) stream beds, sand dunes, orchards, forests (beech, ventrally; metatarsomere 1 as long as metatarsomeres 2+3. podocarp). Mostly open ground; soil moderately moist, Elytra. Subtriangular (contrary to other Harpalini genera). with grass or weeds. Mostly nocturnal; sheltering during Interneurs complete. Rows of setiferous punctures absent the day under logs, stones, plant debris, and in soil bur- on intervals 3, 5, and 7, and on interneur 2. Umbilicate rows. Sometimes active in the sunshine. Biology. setiferous series of interval 9 continuous. Abdomen. Seasonality: September–April, June–July. Tenerals: April. Ventrites 2+3 of male without a setiferous fovea. Ventrites Adult food (in Europe): worms, molluscs, and strawber- 5+6 of both sexes without short setae, with paired ambu- ries; the larva is carnivorous. Occasionally infested with latory setae only. Aedeagus (Fig. 40). Lateral view: strongly mites. Dispersal power. Elytra free along suture. arcuate. Dorsal view: symmetrical (with ostium dorsal, Macropterous. Capable of flight. Moderate runner. Occa- not deflected laterally); dorsal membranous area very wide, sional burrower. Clearly effective as a colonist; currently extending to basal bulb; apical disc present. Internal sac spreading its range both southward and northward. Strongly unarmed. favored by human activities. Collecting techniques. Turn- ing logs and stones, pitfall trapping, using yellow pan traps. Geographic distribution. New Zealand (endemic; North Island). References. Pilgrim, 1963: 837-839 (distribution); Butcher & Emberson, 1981: 60 (distribution, ecology); Johns, 1986: References. Hudson, 1934: 177 (list); Noonan, 1976: 9 31 (distribution, ecology); Lindroth, 1986: 370 (distribu- (taxonomy); Larochelle & Larivière, 2001: 123–124 (cata- tion, ecology, biology, dispersal power); Larochelle & logue). Larivière, 2001: 123 (taxonomy, distribution, ecology, bi- ology, dispersal power) and 2003: 70–71 (ecology, biol- ogy, dispersal power, collecting techniques). Gaioxenus pilipalpis Broun, 1910 E Figures 40, 88, 122, 178; Map p. 147 Gaioxenus pilipalpis Broun, 1910: 8. Lectotype (here desig- Genus Gaioxenus Broun, 1910 E nated): male (BMNH) labelled “Type (circular red-bor- Gaioxenus Broun, 1910: 7. Type species: Gaioxenus pilipalpis dered label; typed) / New Zealand. Broun Coll. 1922-482. Broun, 1910, by monotypy. (typed) / Raurimu Jany. 1909. (hand-written) / (yellow rectangular label without any writing) / Gaioxenus Description. Body length: 8.5–9.0 mm. Boat-shaped (con- pilipalpis (hand-written) / LECTOTYPE Gaioxenus trary to other Harpalini genera). Forebody (head and tho- pilipalpis Broun, 1910 designated by Larochelle & rax) without sparse setiferous micropores dorsally. Head. Larivière 2004 (red label; typed).” Perfect condition. Mandibles moderately long, slightly curved forward, blunt Paralectotypes: 1 male and 2 females (BMNH) bearing apically. Labrum slightly transverse, almost square; apex blue paralectotype labels. curved. Eyes moderately large, convex, widely separated Description. Body length: 8.5–9.0 mm. Slighty convex. from buccal fissure ventrally (by about 1.5× maximum Piceous; lateral margins of pronotum piceous reddish; la- width of antennal scape). Tempora not inflated. Frons with- brum and mandibles reddish; palpi brownish red; anten- out clypeo-ocular prolongations. Antennal pubescence nae, tibiae, and tarsi rusty reddish brown; femora some- starting from middle of antennomere 3. Mentum with a times piceous. Generally glabrous and smooth. tooth medially, moderately shorter than lateral lobes. Men- Microsculpture isodiametric. Dull, without metallic lus- tum and submentum separated by laterally incomplete tre. Head. Moderately large, narrower across eyes than transverse suture. Paraglossae as long as ligula. Palpi with pronotal apex; flat anteriorly, slightly convex posteriorly. last segment fusiform, not truncate apically, with moder- Antennae rather long, reaching basal 1/4 of elytra; antennal ately dense and long pubescence; penultimate segment of scape about 2× longer than its maximum width. Penulti- labial palpi plurisetose on anterior margin. Thorax. mate segment of labial palpi with 2 long setae and 2 short Pronotum transverse; base almost straight, as wide as elytral setae on anterior margin. Thorax. Pronotum (Fig. 122) base; lateral beads complete; anterior bead complete (well very transverse, widest at base; sides converging toward defined medially); posterior bead complete. Scutellum hid- apex, not sinuate; apex concave; lateral depressions widen- 36 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) ing posteriorly; anterior angles moderately developed, ob- Description (Gnathaphanus melbournensis). Body length: tuse-rounded; posterior angles strongly developed, 6.5–7.5 mm. Forebody (head and thorax) without sparse subrectangular; basal foveae shallow, narrow; punctuation setiferous micropores dorsally. Head. Mandibles short, feebly developed. Apex of prosternal lobe with 3–4 long strongly curved forward, blunt apically. Labrum strongly setae and 7–8 short setae. Metepisterna wider than long. transverse; apex slightly emarginate medially. Eyes moder- Elytra. Widest about middle. Shoulders strongly devel- ately large, convex, widely separated from buccal fissure oped, angulate, without a tooth. Subapical sinuations fee- ventrally (by about 1.5× maximum width of antennal scape). ble. Sutural apices angulate. Scutellar striole present. Tempora not inflated. Frons without clypeo-ocular Interneurs shallow, deepening apically, impunctate. Inter- prolongations. Antennal pubescence starting from middle vals impunctate, flat, becoming convex apically. Interval 3 of antennomere 3. Mentum without a tooth medially. with a setiferous puncture behind middle. Aedeagus (Fig. Mentum and submentum separated by complete trans- 40). As for genus. verse suture. Paraglossae longer than ligula. Palpi with last Material examined. 169 specimens, including types segment fusiform, truncate apically, with sparse, moder- (AMNZ, BMNH, CMNZ, ITNZ, JNNZ, LUNZ, NZAC, ately long pubescence; penultimate segment of labial palpi UCNZ). plurisetose on anterior margin. Thorax. Pronotum trans- verse; base slightly emarginate, as wide as elytral base; Geographic distribution (Map p. 147). North Island: lateral beads complete; anterior and posterior beads in- BP, CL, RI, TK, TO, WI, WN, WO. complete medially. Scutellum visible. Apex of prosternal Ecology. Lowland, montane, subalpine, alpine. Mostly lobe pubescent. Legs. Metafemora with 2 long setae on wet forests (beech, broadleaf, podocarp). Also tussock posterior margin. Male protarsi and mesotarsi dilated lat- grasslands. Nocturnal; sheltering during the day often in erally and spongily pubescent ventrally. Segment 4 of burrows dug under stones and logs, also hiding under fallen protarsi and mesotarsi of both sexes without membranous branches and epiphyte crowns. Gregarious. Biology. laminae. Tarsi glabrous dorsally; metatarsomere 5 pubes- Seasonality: September–April. Tenerals: late December– cent (with 6 setae) ventrally; metatarsomere 1 as long as March. Occasionally infested with mites. Dispersal power. metatarsomeres 2+3+4. Elytra. Interneurs complete. Rows Elytra fused along suture. Subapterous. Moderate runnner. of setiferous punctures present on interval 3, absent on Good burrower. Collecting techniques. Turning stones intervals 5 and 7, and on interneur 2. Umbilicate setiferous and logs, pitfall trapping. series of interval 9 continuous. Abdomen. Ventrites 2+3 of References. Larochelle & Larivière, 2001: 123–124 (tax- male without a setiferous fovea. Ventrites 5+6 of both onomy, distribution, ecology, biology, dispersal power). sexes without short setae, with paired ambulatory setae Remarks. Broun described Gaioxenus pilipalpis from 5 only. Aedeagus (Fig. 41). Lateral view: strongly arcuate. specimens, 4 of which could be located in the Natural Dorsal view: asymmetrical (with ostium slightly deflected History Museum, London (BMNH). One of these speci- to the right); dorsal membranous area wide, extending al- mens is designated as lectotype to preserve stability of most to basal bulb; apical disc present. Internal sac armed. nomenclature in the future. This species is unique among Geographic distribution. Australian Region, Oriental New Zealand Harpalini in having a boat-shaped body. In Region, and the Pacific Islands to Japan and New Guinea. the northern part of its range, it occurs at higher elevation, References. Sloane, 1899: 553–554 (key to Australian more or less following the distribution of southern beeches species); Habu, 1973: 56–62 (Japanese taxa); Noonan, 1973: (Nothofagus). 289–290 and 1976: 9 (taxonomy); Moore et al., 1987: 228–232 (synonymy, distribution). Genus Gnathaphanus Macleay, 1825 A Remarks. This genus is in need of revision. Gnathaphanus Macleay, 1825: 20 (originally proposed with subgeneric rank in Trechus Clairville, 1806; first used with generic rank by Chaudoir, 1878: 511). Type spe- cies: Trechus (Gnathaphanus) vulneripennis Macleay, Gnathaphanus melbournensis (Laporte de Castelnau, 1825, by monotypy. A Pachauchenius Macleay, 1864: 116. Type species: 1867) first New Zealand record Pachauchenius laeviceps Macleay, 1864, by monotypy. Figures 41, 89, 123, 179; Map p. 148 Synonymised by Chaudoir, 1878: 511. Harpalus melbournensis Laporte de Castelnau, 1867: 97. Mirosarus Bates, 1878a: 319. Type species: Mirosarus Type locality: Melbourne, Victoria, Australia. insularis Bates, 1878a (=Gnathaphanus melbournensis Harpalus paroensis Laporte de Castelnau, 1867: 98. Type Laporte de Castelnau, 1867), by monotypy. Synonymised locality: Paroo River, New South Wales, Australia. by Sloane, 1899: 553. Synonymised by Chaudoir, 1878: 510. Fauna of New Zealand 53 37

Harpalus marginicollis Laporte de Castelnau, 1867: 103. HB–Waipatiki Beach. South Island: KA–Clarence River Type locality: Melbourne, Victoria, Australia. mouth. Kowhai River mouth. NC–Waiau River. MC–Lin- Synonymised by by Chaudoir, 1878: 510. coln. Mc[C?]lean’s Island. Adventive. Extralimital range: Harpalus adelaideae Laporte de Castelnau, 1867: 108. Type Australia (including Tasmania and Lord Howe Island). First locality: Adelaide and Port Lincoln (South Australia), and King George Sound (Western Australia), Australia. New Zealand record: Mc[?]Clean’s Island, 1985 (PHNZ). Synonymised by by Chaudoir, 1878: 510. Well established. Harpalus planipennis Macleay, 1871: 101. Type locality: Ecology. Lowland. Cultivated fields (carrots), pastures, Gayndah, Queensland, Australia. Synonymised by Sloane, and sand dunes. Open ground; dry sandy soil with sparse 1899: 555. vegetation (grass, weeds). Nocturnal; sheltering during the Harpalus angustatus Macleay, 1871: 102. Type locality: day under logs and stones. Gregarious. Biology. Gayndah, Queensland, Australia. Synonymised by Sloane, 1899: 555. Seasonality: October, January–March. Omnivorous, prob- Harpalus aeneonitens Macleay, 1871: 102. Type locality: ably granivorous (Moore et al., 1987). Dispersal power. Gayndah, Queensland, Australia. Synonymised by by Elytra free along suture. Macropterous. Capable of flight Chaudoir, 1878: 510. (Moore et al., 1987). Moderate runner. Good burrower. Harpalus gayndahensis Macleay, 1871: 102. Type locality: Collecting techniques. Pitfall trapping, turning logs and Gayndah, Queensland, Australia. Synonymised by Sloane, stones. 1899: 555. Harpalus atroviridis Macleay, 1871: 103. Type locality: References. Moore et al., 1987: 229–230 (synonymy, Gayndah, Queensland, Australia. Synonymised by Sloane, distribution, ecology, biology, dispersal power). 1899: 555. Remarks. This introduced species is likely to spread into Mirosarus insularis Bates, 1878a: 319. Type locality: south a wider range of modified habitats. or central Tasmania, Australia. Synonymised by Sloane, 1899: 555. Description. Body length: 6.5–7.5 mm. Slightly convex. Genus Hypharpax Macleay, 1825 N Dark brown; antennal base, palpi, and tibiae yellowish. Hypharpax Macleay, 1825: 22 (originally proposed with Generally glabrous and smooth, except for sparse punc- subgeneric rank in Harpalus Latreille, 1802; first used tuation on pronotum and a series of setiferous punctures with generic rank by Lacordaire, 1854: 282). Type spe- on interval 3. Microsculpture isodiametric, more or less cies: Harpalus (Hypharpax) lateralis Macleay, 1825, by erased on head. Shiny; pronotum and elytra with bronze monotypy. lustre. Head. Moderately large, narrower across eyes than Sagraemerus Redtenbacher, 1868: 13. Type species: Sagraemerus javanus Redtenbacher, 1868, by monotypy. pronotal apex; flat anteriorly, slightly convex posteriorly. Synonymised by Andrewes, 1924: 467. Antennae moderately long, almost reaching pronotal base; antennal scape about 2× longer than its maximum width. Description (New Zealand). Body length: 4.5–7.0 mm. Forebody (head and thorax) without sparse setiferous Penultimate segment of labial palpi with 1–2 long setae and 4–5 short setae on anterior margin. Thorax. Pronotum micropores dorsally. Head. Mandibles short, strongly (Fig. 123) very transverse, widest about middle; sides con- curved forward, blunt apically. Labrum strongly transverse; verging toward base, not sinuate; apex concave; lateral de- apex straight or slightly emarginate medially. Eyes moder- pressions widening posteriorly; anterior angles strongly ately large, convex, widely separated from buccal fissure ventrally (by about 1.5× maximum width of antennal scape). developed, rounded; posterior angles moderately devel- oped, broadly rounded; basal foveae deep, wide; punctua- Tempora not inflated. Frons without clypeo-ocular tion strongly developed (in basal foveae). Apex of prosternal prolongations. Antennal pubescence starting about middle lobe with 2 long setae and several short setae. Metepisterna of antennomere 3. Mentum with a tooth medially, moder- longer than wide. Elytra. Widest behind middle. Shoulders ately shorter than lateral lobes. Mentum and submentum separated by complete transverse suture. Paraglossae as strongly developed, rounded, without a tooth. Subapical sinuations feeble. Sutural apices angulate. Scutellar striole long as ligula. Palpi with last segment fusiform, truncate or present. Interneurs shallow, impunctate. Intervals not apically, with sparse, moderately long pubescence; impunctate, flat. Interval 3 with a series of 5–7 deep, penultimate segment of labial palpi plurisetose or trisetose rather large setiferous punctures. Aedeagus (Fig. 41). As on anterior margin. Thorax. Pronotum transverse, subrectangular; base straight or slightly convex, as wide as for genus. or much narrower than elytral base; lateral beads complete; Material examined. 14 non-type specimens (ITNZ, anterior and posterior beads incomplete medially. Scutellum LUNZ, NZAC, PHNZ, UCNZ). visible. Apex of prosternal lobe pubescent. Legs. Geographical distribution (Map. p. 148). North Island: Metafemora with 4–6 long setae on posterior margin. Male 38 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) protarsi and mesotarsi dilated laterally and spongily pu- Hypharpax antarcticus (Laporte de Castelnau, 1867) E bescent ventrally. Segment 4 of protarsi and mesotarsi of Figures 43, 125, 181; Map p. 148 both sexes without membranous laminae. Tarsi glabrous Harpalus antarcticus Laporte de Castelnau, 1867: 107 dorsally; metatarsomere 5 pubescent (with 4 setae) (redescribed in 1868: 193). Lectotype (here designated): ventrally; metatarsomere 1 subtriangular, short, only about male (MCSN) labelled “N. Zel. Dunedin (hand-written) as long as metatarsomere 2. Elytra. Interneurs complete. / N. Zel. Dunedin (hand-written) Coll. Castelnau (typed) Rows of setiferous punctures absent on intervals 3, 5, and / SYNTYPUS (typed) Harpalus antarcticus Castelnau (hand-written) (red label) / Diaphoromerus antarcticus 7, and on interneur 2. Umbilicate setiferous series of inter- Cast. teste Chaudoir (hand-written) / LECTOTYPE val 9 separated into two major groups, with posterior group Harpalus antarcticus Laporte de Castelnau, 1867 des- continuous. Abdomen. Ventrites 2+3 of male without a ignated by Larochelle & Larivière, 2005 (red label, setiferous fovea. Ventrites 5+6 of both sexes without short typed).” Excellent condition. setae, with paired ambulatory setae only. Aedeagus. Lat- Hypharpax antarcticus: Bates, 1874: 272. eral view: strongly arcuate. Dorsal view: asymmetrical (with Diaphoromerus antarcticus: Chaudoir, 1878: 485. ostium deflected to the right); dorsal membranous area Hypharpax antarcticus: Hudson, 1934: 37, 177. wide, extending to basal bulb; apical disc present. Internal Description. Body length: 4.5–5.5 mm. Slightly convex. sac armed. Green or piceous black; base of antennae (segments 1+2) Geographic distribution. Australia (including Tasmania and tibiae (2/3rd) pale, reddish brown. Generally glabrous and Lord Howe Island), New Guinea, Indonesia, New Zea- and smooth. Microsculpture moderately transverse. Some- land. what dull, with aeneous or bronze metallic lustre. Head. Moderately large, narrower across eyes than pronotal apex; References. Hudson, 1934: 177 (list); Noonan, 1973: 288– flat anteriorly, slightly convex posteriorly. Antennae mod- 289 and 1976: 9 (taxonomy); Moore et al., 1987: 237–240 erately long, reaching about pronotal base; antennal scape (list of Australian species); Moore, 1992: 168 (distribu- about 2× longer than its maximum width; antennomeres 8– tion); Larochelle & Larivière, 2001: 124–125 (catalogue). 10 almost square. Palpi not narrowly truncate apically; Remarks. This genus is in need of revision. penultimate segment of labial palpi plurisetose (with 2 long setae and 2–3 short setae) on anterior margin. Tho- rax. Pronotum (Fig. 125) very transverse, subrectangular, widest about middle; base straight, as wide as elytral base; Key to species of Hypharpax sides converging toward base, moderately convex, not 1 Pronotum (Fig. 125): sides moderately convex; posterior sinuate; apex straight; lateral depressions absent; anterior angles obtuse; base straight, as wide as elytral base; angles feebly developed, rounded; posterior angles strongly basal foveae shallow, wide; base finely punctate. developed, obtuse; basal foveae shallow, wide; punctua- Antennomeres 8–10 almost square. Elytra (Fig. 181) tion feebly developed (basally). Apex of prosternal lobe shorter (about 2.3× longer than wide). Penultimate with 3–4 long setae and numerous short setae. Metepisterna segment of labial palpi plurisetose (Fig. 9). Aedeagus longer than wide. Elytra. Widest about middle; approxi- (Fig. 43): apex swollen, button-like. Body length usually mately 2.3× longer than wide. Shoulders strongly devel- less than 5.5 mm. [South Island] ...... oped, angulate, without a tooth. Subapical sinuations fee- ...... (p. 38)... antarcticus (Laporte de Castelnau) ble. Sutural apices angulate-rounded. Scutellar striole ab- — Pronotum (Fig. 124): sides strongly convex; posterior sent or present. Interneurs shallow, impunctate. Intervals angles broadly rounded; base slightly convex, much impunctate, flat. Interval 3 with a setiferous puncture be- narrower than elytral base; basal foveae deep, rather hind middle. Aedeagus (Fig. 43). Lateral view: as for ge- narrow; base coarsely punctate. Antennomeres 8–10 nus; extremity of apex swollen, button-like. Dorsal view: elongate. Elytra (Fig. 180) longer (about 2.8× longer as for genus; ostium strongly deflected to the right. than wide). Penultimate segment of labial palpi trisetose Material examined. 108 specimens, including types (Fig. 10). Aedeagus (Fig. 42): apex barely inflated, not (AMNZ, CMNZ, ITNZ, LUNZ, MCSN, NZAC, PHNZ, button-like. Body length usually over 5.5 mm. [North UCNZ). Island, South Island, Chatham Islands] ...... Geographic distribution (Map p. 148). South Island: ...... (p. 39)... australis (Dejean) CO, DN, KA, MB, MC, MK, NC, NN, OL, SC. Offshore Islands: CH. Ecology. Lowland, montane, subalpine, alpine. Tussock grasslands, cultivated fields (lucerne, Festuca), pastures, Fauna of New Zealand 53 39 and gardens. Also river banks. Open ground; soil covered with aeneous or bronze metallic lustre. Head. Moderately with grass or weeds. Mostly diurnal; usually active in the large, narrower across eyes than pronotal apex; flat sunshine; sheltering on cloudy days under stones and at anteriorly, slightly convex posteriorly. Antennae moder- the base of tussock clumps. Biology. Seasonality: through- ately long, reaching about pronotal base; antennal scape out the year. Predators: starlings. Dispersal power. Elytra about 2× longer than its maximum width; antennomeres 8– free along suture. Macropterous. Frequent flier. Moderate 10 elongate. Palpi narrowly truncate apically; penultimate runner. Regular climber (on plants). Strongly favoured by segment of labial palpi trisetose (with 2 long setae and 1 human activities. Collecting techniques. Pitfall trapping, short seta) on anterior margin. Thorax. Pronotum (Fig. turning stones, sweeping plants. 124) very transverse, subrectangular, widest before mid- References. Hudson, 1934: 37 (distribution, ecology); dle; base slightly convex, much narrower than elytral base; Noonan, 1973: 289 (taxonomy); Johns, 1986: 31 (distribu- sides converging toward base (more so than in antarcticus), tion); Larochelle & Larivière, 2001: 124 (taxonomy, distri- strongly convex, not sinuate; apex straight; lateral depres- bution, ecology, biology, dispersal power). sions absent; anterior angles feebly developed, rounded; posterior angles moderately developed, broadly rounded; Remarks. Twelve syntypes of Harpalus antarcticus were basal foveae deep, rather narrow; punctuation strongly obtained from the Castelnau collection in Genova (MCSN), developed (in basal foveae and basally). Apex of prosternal one of which (a male collected from Dunedin) is here lobe with 3–4 long setae and numerous short setae. designated as a lectotype to preserve stability of nomen- Metepisterna longer than wide. Elytra. Widest about mid- clature in the future. This male is the middle specimen of dle; approximately 2.8× longer than wide. Shoulders a series of 3 syntypes glued to the same card. strongly developed, angulate, without a tooth. Subapical Although morphologically close, H. antarcticus and sinuations feeble. Sutural apices angulate-rounded. Scutellar H. australis can be easily diagnosed based on external striole present. Interneurs shallow, impunctate. Intervals features and male aedeagus. In addition, H. antarcticus impunctate, flat. Interval 3 with a setiferous puncture be- does not occur in the North Island. hind middle. Aedeagus (Fig. 42). Lateral view: as for ge- nus; extremity of apex barely inflated dorsally. Dorsal view: Hypharpax australis (Dejean, 1829) A as for genus; ostium slightly deflected to the right. Figures 42, 90, 124, 180; Map p. 148 Material examined. 502 specimens, including H. Harpalus australis Dejean, 1829: 385. Type locality: Aus- abstrusus type (AMNZ, BBNZ, ITNZ, JNNZ, LUNZ, tralia (as Nouvelle-Hollande). MNHN, MONZ, NZAC, OMNZ, PHNZ, UCNZ). Harpalus inornatus Germar, 1848: 169. Type locality: Ad- elaide, South Australia. Synonymised by Chaudoir, 1878: Geographic distribution (Map p. 148). North Island: 484. AK, BP, CL, GB, HB, ND, RI, TK, TO, WI, WN, WO. Harpalus coxii Laporte de Castelnau, 1867: 107 (redescribed South Island: CO, KA, MB, MC, MK, NN, OL, SC, SD. in 1868: 193). Type locality: Clarence River, New South Offshore Islands: CH, TH. Adventive. Extralimital range: Wales, Australia. Synonymised by Chaudoir, 1878: 484. Australia (including Tasmania and Lord Howe Island). First Hypharpax australis: Bates, 1874: 272. New Zealand records: Mt Albert, AK, 1916 (NZAC); New Diaphoromerus australis: Chaudoir, 1878: 484. Zealand (Redtenbacher, 1868: 15), Auckland, AK (MNHN, Hypharpax australis: Broun, 1880: 51. Hypharpax abstrusus Bates, 1878c: 23. Lectotype (here des- as H. abstrusus Bates, 1878). Well established. ignated): male (MNHN) labelled “Auckland N.Z. (hand- Ecology. Lowland, montane. Sand dunes, pastures, culti- written) / Coll. Chaudoir (hand-written) / Ex Musaeo vated fields (lucerne, carrot, pea), farmlands, gardens, tus- H.W. BATES 1892 (typed) / Hypharpax abstrusus Bates sock grasslands, river banks, scrublands. Open ground; (hand-written) / LECTOTYPE Hypharpax abstrusus sandy or clayey soil covered with grass or weeds. Mostly Bates, 1878 designated by Larochelle & Larivière 2004 (red label; typed).” Perfect condition. New synonym. diurnal; usually active in the sunshine; sheltering on cloudy Hypharpax parvus Chaudoir, 1878: 500. Type locality: days in burrows dug at the base of plants, under stones, Southern Australia. Synonymised by Moore, in Moore et pieces of wood, logs, dead leaves, Muehlenbeckia-clumps, al., 1987: 237. and at the base of Spinifex plants. Gregarious. Biology. Hypharpax (Harpalus) australis: Blackburn, 1892: 83. Seasonality: throughout the year, except May–June. Hypharpax australis: Hutton, 1904: 351. Tenerals: February. Omnivorous, probably granivorous Description. Body length: 5.5–7.0 mm. Slightly convex. (Moore et al., 1987). Predators: starlings. Regularly in- Green or piceous black; base of antennae (segments 1+2) fested with mites. Dispersal power. Elytra free along su- and tibiae (2/3rd) pale, reddish brown. Generally glabrous ture. Macropterous. Frequent flier. Moderate runner. Oc- and smooth. Microsculpture isodiametric. Somewhat dull casional climber (on shrubs and plants). Clearly effective 40 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) as a colonist. Strongly favoured by human activities. Col- long setae on posterior margin. Male protarsi and mesotarsi lecting techniques. Pitfall trapping, digging at the base of dilated laterally and spongily pubescent ventrally. Seg- plants, turning debris, light trapping. ment 4 of protarsi and mesotarsi of both sexes without References. Walker, 1904: 76 (as abtrusus; distribution, membranous laminae. Tarsi pubescent (with numerous se- ecology); Thomson, 1922: 284 (distribution); Pilgrim, 1963: tae) dorsally; metatarsomere 5 pubescent (with numerous 842 (distribution); Noonan, 1973: 289 (taxonomy); Johns, setae) ventrally; metatarsomere 1 as long as metatarsomeres 1980: 63 (as abtrusus; distribution, ecology); Moeed, 1980: 2+3. Elytra. Interneurs complete. Rows of setiferous punc- 250 (biology); Butcher & Emberson, 1981: 64 (distribu- tures absent on intervals 3, 5, and 7, and on interneur 2. tion, ecology); Johns, 1986: 31 (distribution, ecology); Umbilicate setiferous series of interval 9 continuous. Ab- Moore et al., 1987: 237–238 (synonymy, distribution, ecol- domen. Ventrites 2+3 of male without a setiferous fovea. ogy, biology, dispersal power); Kuschel, 1990: 24, 40 (dis- Ventrites 5+6 of both sexes without short setae, with paired tribution, ecology, biology, dispersal power); Moore, 1992: ambulatory setae only. Aedeagus (Fig. 44). Lateral view: 168 (distribution); Patrick et al., 1992: 272 (distribution); slightly arcuate. Dorsal view: symmetrical (with ostium Townsend, 1997: 16 and 1998:18, 21 (distribution); dorsal, not deflected laterally); dorsal membranous area Emberson, 1998: 30 (distribution, ecology, biology); very wide, extending almost to basal bulb; apical disc ab- Larochelle & Larivière, 2001: 125 (taxonomy, distribution, sent. Internal sac unarmed. ecology, biology, dispersal power). Geographic distribution. New Zealand (endemic; Three Remarks. Bates’ original description was based on “a sin- Kings Islands). gle pair [male symbol, female symbol]” from “Auckland.” Remarks. The generic name is derived from Maori (the The male syntype was obtained from the Bates’ collection Polynesian people who colonised New Zealand) and in Paris (MNHN); it is here selected as lectotype to pre- Harpalus (the type genus of the tribe Harpalini). This serve stability of nomenclature in the future. The wherea- very distinctive monotypic taxon is characterised by long bouts of the female syntype are unknown. This species mandibles and antennal scapes, a strongly emarginate la- occurs on both main islands of New Zealand. See also brum, relatively small eyes, and a suborbicular pronotum. Remarks under H. antarcticus.

Maoriharpalus sutherlandi new species E Genus Maoriharpalus new genus E Figures 44, 91, 126, 182; Map p. 148 Type species. Maoriharpalus sutherlandi new species, by Maoriharpalus sutherlandi Larochelle & Larivière, new spe- present designation. cies. Holotype: male (NZAC) labelled “Three Kings Is Great I. Nov. 70 NZ. Ent. Div. Exp. (typed) / G. Ramsay Description. Body length: 12.0–13.0 mm. Forebody (head (typed) / Castaway Camp (typed) / HOLOTYPE [male and thorax) without sparse setiferous micropores dorsally. symbol] Maoriharpalus sutherlandi Larochelle & Head. Mandibles very long (about 6x their maximum Larivière, 2004 (red label; typed).” Paratypes: 3 males width), slightly curved forward, blunt apically. Labrum (2 NZAC, 1 AMNZ) from Great Island, bearing blue strongly transverse; apex strongly emarginate medially. paratype labels. Eyes moderately large, convex, widely separated from buccal Description. Body length: 12.0–13.0 mm. Slightly con- fissure ventrally (by about 2× maximum width of antennal vex. Piceous black; antennae (except segment 1), palpi, and scape). Tempora not inflated. Frons without clypeo-ocu- tarsi rufous. Generally glabrous and smooth. lar prolongations. Antennal pubescence starting from basal Microsculpture isodiametric on head, moderately trans- third of antennomere 3; antennal scape very long (about 5- verse on pronotum, very transverse (with microlines) on 6x longer than maximum width; contrary to other elytra. Shiny, without metallic lustre; elytra iridescent. Anisodactylina genera). Mentum without tooth medially. Head. Moderately large, narrower across eyes than pronotal Mentum and submentum separated by complete trans- apex; excavated anteriorly, flat posteriorly. Antennae very verse suture. Paraglossae longer than ligula. Palpi with last long, reaching middle of elytra; antennal scape elongate, segment cylindrical, truncate apically, with moderately about 5–6× longer than its maximum width. Penultimate dense, long pubescence; penultimate segment of labial palpi segment of labial palpi with 5–6 long setae on anterior plurisetose on anterior margin. Thorax. Pronotum margin. Thorax. Pronotum (Fig. 126) suborbicular, widest suborbicular; base strongly convex, moderately narrower before middle; sides converging toward base, not sinuate; than elytral base; lateral beads complete; anterior and pos- apex convex; lateral depressions widening posteriorly; an- terior beads incomplete medially. Scutellum visible. Apex terior angles strongly developed, obtuse; posterior angles of prosternal lobe pubescent. Legs. Metafemora with 3 feebly developed, broadly rounded; basal foveae deep, nar- Fauna of New Zealand 53 41 row; punctuation feebly developed. Apex of prosternal spongily pubescent ventrally. Segment 4 of protarsi and lobe with 4-6 long setae. Metepisterna wider than long. mesotarsi of both sexes without membranous laminae. Tarsi Elytra. Widest about middle. Shoulders well developed, glabrous dorsally; metatarsomere 5 pubescent (with 4 se- rounded, without a tooth. Subapical sinuations feeble. tae) ventrally; metatarsomere 1 parallel-sided, very long, Sutural apices angulate. Scutellar striole present. Interneurs almost as long as metatarsomeres 2+3+4. Elytra. Interneurs shallow, deepening apically, impunctate. Intervals complete. Rows of setiferous punctures absent on inter- impunctate, flat, becoming convex apically. Interval 3 with- vals 3, 5, and 7, and on interneur 2. Umbilicate setiferous out setiferous puncture behind middle. Aedeagus (Fig. series of interval 9 separated into two major groups, with 44). As for genus. posterior group continuous. Abdomen. Ventrites 2+3 of Material examined. 10 specimens, including types male without a setiferous fovea. Ventrites 5+6 of both (AMNZ, NZAC). sexes without short setae, with paired ambulatory setae only. Aedeagus. No male seen. Geographic distribution (Map p. 148). Offshore Islands: TH-Great Island. South West Island. Geographic distribution. Australian Region, Neartic Re- gion, Neotropical Region. Ecology. Lowland. Wet forests (broadleaf, podocarp). Shaded ground. Nocturnal; sheltering during the day under References. Noonan, 1973: 293–338 and 1976: 10–11 stones. Biology. Seasonality: October–November. Tenerals: (taxonomy). Moore et al., 1987: 232–237 (distribution). November. Dispersal power. Elytra fused along suture. Remarks. Australian representatives of this genus are in Subapterous. Moderate runner. Collecting need of revision. techniques.Turning stones.

Remarks. This species is named after our friend O. R. W. A Sutherland (former Science Manager, Landcare Research) Subgenus Anisotarsus Chaudoir, 1837 for his special help and encouragement in establishing our Anisotarsus Chaudoir, 1837: 41. Type species: Anisotarsus brevicollis Chaudoir, 1837, designated by Emden, 1953: new life and career in New Zealand. The mouthparts indi- 519. cate that the species may feed on hard-bodied inverte- Diaphoromerus Chaudoir, 1843: 402. Type species: brates like snails. Diaphoromerus iridipennis Chaudoir, 1843, by monotypy. Eurytrichus LeConte, 1848: 387. Type species: Feronia Genus Notiobia Perty, 1830 A terminata Say, 1823, designated by Emden, 1953: 525. Notiobia Perty, 1830: 13. Type species: Notiobia nebrioides Stilbolidus Casey, 1914: 171. Type species: Harpalus Perty, 1830, by monotypy. mexicanus Dejean, 1829, by original designation. Description (Notiobia quadricollis). Body length: about Geographic distribution. As for genus. 8.0 mm. Forebody (head and thorax) without sparse References. As for genus. setiferous micropores dorsally. Head. Mandibles moder- ately long, slightly curved forward, blunt apically. Labrum strongly transverse; apex slightly emarginate medially. Eyes Notiobia (A.) quadricollis (Chaudoir, 1878) A first New moderately large, convex, widely separated from buccal Zealand record fissure ventrally (by about 2× maximum width of antennal Figures 92, 127, 183; Map p. 148 scape). Tempora not inflated. Frons without clypeo-ocu- Diaphoromerus quadricollis Chaudoir, 1878: 86. Type lar prolongations. Antennal pubescence starting from mid- locality: northern Australia. dle of antennomere 3. Mentum with a tooth medially, mod- erately shorter than lateral lobes. Mentum and submentum Notiobia (A.) quadricollis: Noonan, 1976: 10. separated by complete transverse suture. Paraglossae as Description. Body length: 8.2 mm. Moderately convex. long as ligula. Palpi with last segment fusiform, truncate Black; antennomere 1 rufous. Generally glabrous and apically, with sparse, moderately long pubescence; penul- smooth. Microsculpture isodiametric. Dull, without me- timate segment of labial palpi plurisetose on anterior mar- tallic lustre. Head. Moderately large, narrower across eyes gin. Thorax. Pronotum transverse, subrectangular; base than pronotal apex; flat anteriorly, slightly convex straight, as wide as elytral base; lateral beads complete; posteriorly. Antennae moderately long, reaching pronotal anterior bead incomplete medially; posterior bead com- base; antennal scape about 2× longer than its maximum plete. Scutellum visible. Apex of prosternal lobe pubes- width. Penultimate segment of labial palpi with 2 long cent. Legs. Metafemora with 2 long setae on posterior setae and 6 short setae on anterior margin. Thorax. margin. Male protarsi and mesotarsi dilated laterally and Pronotum (Fig. 127) very transverse, widest about middle; 42 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) sides converging toward base, not sinuate; apex straight; Scutellum visible. Apex of prosternal lobe pubescent. Legs. lateral depressions widening posteriorly; anterior angles Metafemora with 2 long setae on posterior margin. Male moderately developed, obtuse; posterior angles strongly protarsi dilated laterally and spongily pubescent ventrally. developed, subrectangular; basal foveae deep, moderately Male mesotarsi dilated laterally and spongily pubescent wide; punctuation strongly developed (in basal foveae). ventrally (atratus) or unmodified (hoarei, lesagei). Seg- Apex of prosternal lobe with 3–4 long setae. Metepisterna ment 4 of protarsi and mesotarsi of both sexes without longer than wide. Elytra. Widest about middle. Shoulders membranous laminae. Tarsi pubescent (with numerous se- well developed, angulate, with a tooth. Subapical sinuations tae) dorsally; metatarsomere 5 pubescent (with numerous moderate. Sutural apices angulate. Scutellar striole present. setae) ventrally; metatarsomere 1 as long as metatarsomeres Interneurs shallow, impunctate. Intervals impunctate, flat. 2+3. Elytra. Interneurs complete. Rows of setiferous punc- Interval 3 with a setiferous puncture behind middle. tures absent on intervals 3, 5, and 7, and on interneur 2. Aedeagus. No male seen. Umbilicate setiferous series of interval 9 continuous. Ab- Material examined. 1 non-type specimen (NZAC). domen. Ventrites 2+3 of male without a setiferous fovea. Ventrites 5+6 of both sexes without short setae, with paired Geographic distribution (Map p. 148). North Island: ambulatory setae only. Aedeagus. Lateral view: strongly ND–Spirits Bay (1957). Adventive. Extralimital distribu- arcuate. Dorsal view: symmetrical (with ostium dorsal, tion: Australia (mainland). Doubtfully established. not deflected laterally) or asymmetrical (with ostium de- Ecology. One dead specimen found on a beach. Biology. flected slightly to the right); dorsal membranous area wide, Seasonality: unknown. Omnivorous, probably granivorous extending to basal bulb or almost; apical disc present. In- (Moore et al., 1987). Dispersal power. Elytra free along ternal sac armed or unarmed. suture. Macropterous. Capable of flight (Moore et al., Geographic distribution. New Zealand (endemic; North 1987). Moderate runner (after leg morphology). Island). References. Moore et al., 1987: 236 (synonymy, distri- References. Hudson, 1934: 174 (list); Britton, 1964b: bution, ecology, biology, dispersal power). 522–523, 526 (taxonomy); Noonan, 1976: 9 (taxonomy); Remarks. Since no additional specimen of this species has Larochelle & Larivière, 2001: 125 (catalogue). been found following the discovery of a single specimen in Remarks. Although P. hoarei appears to be less closely 1957, it seems unlikely that natural populations of this related to P. atratus and P. lesagei than they are to each species have established themselves in New Zealand. other on the basis of morphology, the shared morphologi- cal characters defining the genus and the stability of these Genus Parabaris Broun, 1881 E characters within each species, suggest that they share a Parabaris Broun, 1881: 654. Type species: Parabaris atratus common ancestor. Broun, 1881, by monotypy. Description. Body length: 9.5–20.0 mm. Forebody (head Key to species of Parabaris and thorax) without sparse setiferous micropores dorsally. Head. Mandibles moderately long, slightly curved for- 1 Pronotum (Fig. 130): base emarginate; sides convex, not ward, blunt apically. Labrum strongly transverse; apex sinuate. Elytra very iridescent; interval 3 with 2 slightly emarginate medially. Eyes moderately large and setiferous punctures subapically. Body dark brown, convex, widely separated from buccal fissure ventrally (by length 10.5 mm or less ...(p. 44)... hoarei new species 2–3× maximum width of antennal scape). Tempora not — Pronotum (Fig. 128–129): base straight; sides sinuate. inflated. Frons without clypeo-ocular prolongations. Elytra moderately iridescent; interval 3 without Antennal pubescence starting from basal 1/3 of antennomere setiferous puncture subapically. Body black, length 3. Mentum with a tooth medially, moderately shorter than 15.5 mm or more ...... 2 lateral lobes. Mentum and submentum separated by com- 2(1) Head narrower across eyes than pronotal apex (Fig. plete transverse suture. Paraglossae longer than ligula. Palpi 184). Pronotum (Fig. 128) very transverse. [Body stout, with last segment fusiform or rather cylindrical, truncate length 16.0–20.0 mm] ...... (p. 43)... atratus Broun or not apically, with moderately dense and long pubes- — Head as wide across eyes as pronotal apex (Fig. 185). cence; penultimate segment of labial palpi plurisetose on Pronotum (Fig. 129) moderately transverse. [Body anterior margin. Thorax. Pronotum transverse; base straight slender, length 15.5–16.5 mm] ...... (atratus, lesagei) or emarginate, as wide as or narrower ...... (p. 43)... lesagei new species than elytral base; lateral beads complete; anterior bead in- complete medially and ill-defined; posterior bead complete. Fauna of New Zealand 53 43

Parabaris atratus Broun, 1881 E Ecology. Lowland, montane. Wet or moist forests Figures 45, 93, 128, 184; Map p. 148 (broadleaf, podocarp, beech) and swamp forests: along Parabaris atratus Broun, 1881: 655. Holotype: female mud flats, gullies, and streams. Shaded ground; soil cov- (BMNH) labelled “Type (circular red-bordered label; ered with dead leaves. Nocturnal; sheltering during the day typed) / 1144. (typed) / New Zealand. Broun Coll. Brit. under logs (mostly) and stones, and in rotten logs. Biol- Mus. 1922-482. (typed) / Parua. Whangarei (hand-writ- ogy. Seasonality: September–June, August. Tenerals: Janu- ten) / Parabaris atratus. [female symbol] (hand-written).” ary, April (mostly). Occasionally infested with mites and Poor condition; antennae and legs missing some antennomeres and tarsomeres; forebody (head and tho- fungi (Laboulbeniales). Dispersal power. Elytra fused rax) separated from hindbody. along suture. Subapterous. Moderate runner. Collecting techniques. Pitfall trapping, turning logs and stones, us- Description. Body stout (compared to lesagei), length: ing yellow pan traps. 16.0–20.0 mm. Slightly convex. Black; legs piceous black; antennae (except segment 1), palpi, and tarsi rufous. Gen- Reference. Larochelle & Larivière, 2001: 125 (taxonomy, erally glabrous and smooth. Microsculpture isodiametric distribution, ecology, biology, dispersal power). on head, moderately transverse on thorax, very transverse Remarks. This species and P. lesagei are closer morpho- (with microlines) on elytra. Shiny, without metallic lustre. logically than either is to P. hoarei. Elytra moderately iridescent. Head. Moderately large, nar- rower across eyes than pronotal apex; flat anteriorly, slightly convex posteriorly. Eyes widely separated from buccal Parabaris lesagei new species E × fissure ventrally (by about 2 maximum width of antennal Figures 46, 129, 185; Map p. 148 scape). Antennae moderately long, reaching pronotal base; Parabaris lesagei Larochelle & Larivière, new species. antennal scape elongate, about 4× longer than its maximum Holotype: male (NZAC) labelled “NEW ZEALAND WN width. Palpi cylindrical, slightly truncate apically; penul- Kaitoke Regional Park (Waterworks Rd end) 300m timate segment of labial palpi with 4–6 long setae on ante- 26.XI.1996 Larochelle, Larivière (typed) / Wet broadleaf rior margin. Thorax. Pronotum (Fig. 128) very transverse, forest Along stream banks, under stones. (typed) / widest before middle; sides converging toward base, feebly HOLOTYPE [male symbol] Parabaris lesagei Larochelle sinuate; base straight, slightly narrower than elytral base & Larivière, 2004 (red label; typed).” Paratypes: 2 males (1 NZAC, 1 MONZ), 1 female (NZAC) from the same (less so than in lesagei); apex concave; lateral beads widen- locality as the holotype, bearing blue paratype labels. ing about middle (as in lesagei); lateral depressions widen- Description. Body slender (compared to atratus), length: ing posteriorly; anterior angles strongly developed, acutely 15.5–16.5 mm. Slightly convex. Black; legs piceous black; rounded; posterior angles strongly developed, rectangular; basal foveae deep, wide; punctuation feebly developed. antennae (except segment 1), palpi, and tarsi rufous. Gen- erally glabrous and smooth. Microsculpture isodiametric Apex of prosternal lobe with 8–11 long setae. Metepisterna on head, moderately transverse on thorax, and very trans- as wide as long. Legs. Male mesotarsi dilated laterally and verse (with microlines) on elytra. Shiny, without metallic spongily pubescent ventrally. Elytra. Widest before mid- lustre. Elytra moderately iridescent. Head. Moderately dle. Shoulders strongly developed, angulate, with a tooth. Subapical sinuations strong. Sutural apices angulate. large, as wide across eyes as pronotal apex; flat anteriorly, slightly convex posteriorly. Eyes widely separated from Scutellar striole present. Interneurs moderately deep, buccal fissure ventrally (by about 3× maximum width of impunctate. Intervals impunctate, rather flat in basal half, antennal scape). Antennae moderately long, reaching slightly convex in apical half. Interval 3 without setiferous pronotal base; antennal scape elongate, about 4× longer puncture behind middle. Aedeagus (Fig. 45). Lateral view: strongly arcuate (including apical 1/2), thinner than in other than its maximum width. Palpi cylindrical, slightly trun- cate apically; penultimate segment of labial palpi with 4-6 Parabaris species; apex narrowly pointed. Dorsal view: long setae on anterior margin. Thorax. Pronotum (Fig. asymmetrical (with ostium slightly deflected to the right); 129) moderately transverse (less than in atratus), widest dorsal membranous area very wide, extending to basal bulb; before middle; sides converging toward base, moderately apical disc present, rounded-triangular. Internal sac un- armed. sinuate; base straight, much narrower than elytral base (more so than in atratus); apex concave; lateral beads wid- Material examined. 148 specimens, including type ening about middle (as in atratus); lateral depressions widen- (AMNZ, BBNZ, BMNH, CMNZ, ITNZ, JNNZ, LUNZ, ing posteriorly; anterior angles strongly developed, acutely MONZ, NZAC, UCNZ). rounded; posterior angles strongly developed, rectangular; Geographic distribution (Map p. 148). North Island: basal foveae deep, wide; punctuation feebly developed. AK, BP, CL, GB, ND, TK, WO. Apex of prosternal lobe with 8–11 long setae. Metepisterna 44 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) as wide as long. Legs. Male mesotarsi unmodified, neither very iridescent. Head. Moderately large, narrower across dilated laterally nor spongily pubescent ventrally. Elytra. eyes than pronotal apex; flat anteriorly, slightly convex Widest before middle (as in atratus). Shoulders strongly posteriorly. Eyes widely separated from buccal fissure developed, angulate, with a tooth. Subapical sinuations ventrally (by about 2× maximum width of antennal scape). strong. Sutural apices rounded. Scutellar striole present. Antennae moderately long, reaching about pronotal base; Interneurs moderately deep, impunctate. Intervals antennal scape about 2× longer than its maximum width. impunctate, flat in basal half, slightly convex in apical half. Palpi fusiform, not truncate apically; penultimate segment Interval 3 without setiferous puncture behind middle. of labial palpi with 2–4 long setae and 1–3 short setae on Aedeagus (Fig. 46). Lateral view: strongly arcuate (with anterior margin. Thorax. Pronotum (Fig. 130) very trans- apical half rather straight); apex narrowly pointed, with verse, widest about middle; sides converging toward base, apical disc partially visible dorsally. Dorsal view: asym- convex; base emarginate, slightly narrower than elytral base; metrical (with ostium slightly deflected to the right); dor- apex slightly concave; lateral beads not widening about sal membranous area moderately wide, extending almost to middle; lateral depressions widening posteriorly; anterior basal bulb; apical disc present, rectangular. Internal sac angles moderately developed, obtuse; posterior angles armed. moderately developed, rounded; basal foveae deep, wide; Material examined. 11 specimens, including types punctuation feebly developed. Apex of prosternal lobe (AMNZ, ITNZ, JNNZ, LUNZ, MONZ, NZAC). with usually 3-4 long setae and several short setae. Metepisterna wider than long. Legs. Mesotarsi unmodi- Geographic distribution (Map p. 148). North Island: fied, neither dilated laterally nor spongily pubescent BP–Te Koau. TK–Mount Messenger. WN–Belmont. ventrally. Elytra. Widest about middle. Shoulders strongly Kaikoke Regional Park. Orongorongo Valley. Otaki Forks. developed, angulate, without a tooth. Subapical sinuations Rimutaka Hill. Rimutaka Forest Park, Junction of feeble. Sutural apices angulate. Scutellar striole absent. Orongorongo River and Turere Stream. Interneurs moderately deep, impunctate. Intervals Ecology. Lowland, montane. Wet forests (broadleaf, beech): impunctate, slightly convex. Interval 3 with two setiferous along streams and gullies. Shaded ground; soil covered with punctures subapically. Aedeagus (Fig. 47). Lateral view: dead leaves. Nocturnal; sheltering during the day under big strongly arcuate (less so in apical half); apex abruptly nar- stones and logs. Biology. Seasonality: November–Febru- rowed. Dorsal view: symmetrical (with ostium dorsal, not ary. Tenerals: March. Dispersal power. Elytra fused along deflected laterally); dorsal membranous area moderately suture. Subapterous. Moderate runner. Good burrower. wide, extending to basal bulb; apical disc present, rounded- Collecting techniques. Turning big stones and logs, pit- triangular (shorter than in atratus). Internal sac armed. fall trapping. Material examined. 15 specimens, including types Remarks. This species is named after our close friend (MONZ, NZAC, UCNZ). Laurent LeSage (Agriculture and Agri-Food Canada, Ot- Geographic distribution (Map p. 148). North Island: tawa, Canada) for his special help and encouragement in ND–Kapowairua. Spirits Bay. Tom Bowling Bay. our life and career. See also Remarks under P. atratus. Unuwhao. Waipuna Stream. Ecology. Lowland. Wet forests (broadleaf). Shaded ground. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: November–January, August. E Parabaris hoarei new species Tenerals: November, early January. Dispersal power. Figures 47, 94, 130, 186; Map p. 148 Elytra fused along suture. Subapterous. Moderate runner. Parabaris hoarei Larochelle & Larivière, new species. Collecting techniques. Turning stones and logs. Holotype: male (NZAC) labelled “UNUWHAO SPIR- ITS BAY 11.I. 57. R.A. CUMBER. (hand-written) / Remarks. The presence of 2 setiferous punctures subapi- HOLOTYPE [male symbol] Parabaris hoarei Larochelle cally on the elytral interval 3, the body shape, brownish & Larivière, 2004 (red label; typed).” Paratypes: 3 fe- colour, and very iridescent elytra set this taxon apart from males (2 NZAC, 1 MONZ) from the same locality as the its congeners. This is the only Parabaris species so far holotype, bearing blue paratype labels. recorded from and restricted to the tip of the Aupouri Description. Body length: 9.5–10.5 mm. Slightly convex. Peninsula. This species is named after our friend and col- Piceous brown; antennae, palpi, and legs rufous. Generally league R. J. B. Hoare (Landcare Research, Auckland) for glabrous and smooth. Microsculpture moderately trans- his special help and encouragement in our entomological verse on head and pronotum, very transverse (with studies and for his special talent and dedication as a manu- microlines) on elytra. Shiny, without metallic lustre. Elytra script reviewer. Fauna of New Zealand 53 45

Genus Triplosarus Bates, 1874 E Triplosarus novaezelandiae (Laporte de Castelnau, Triplosarus Bates 1874: 270 (redescribed in 1875: 308). Type 1867) E species: Triplosarus fulvescens Bates, 1874 (=Harpalus Figures 48, 95, 131, 187; Map p. 148 novaezelandiae Laporte de Castelnau, 1867), by Harpalus novaezelandiae Laporte de Castelnau, 1867: 108 monotypy. (redescribed in 1868: 194). Lectotype (here designated): Description. Body length: 7.5–10.0 mm. Forebody (head male (MCSN) labelled “Auck[land] [N. Zelanda] (hand- and thorax) without sparse setiferous micropores dorsally. written) Coll. Castelnau (typed) / Harp. novae zelan- Head. Mandibles moderately long, strongly curved for- diae. auck. (hand-written)/ Harpalus Novae Zelandiae ward, blunt apically. Labrum strongly transverse; apex Cast. (hand-written) / SYNTYPUS (typed) Harpalus novae zelandiae Castelnau (hand-written) (red label) / straight or slightly emarginate medially. Eyes moderately LECTOTYPE Harpalus novaezelandiae Laporte de large, convex, widely separated from buccal fissure Castelnau, 1867 designated by Larochelle & Larivière, × ventrally (by about 1.3 maximum width of antennal scape). 2005 (red label, typed).” Excellent condition. Tempora not inflated. Frons without clypeo-ocular Triplosarus fulvescens Bates, 1874: 271 (redescribed in 1875: prolongations. Antennal pubescence starting from middle 309). Lectotype (here designated): male (MNHN) la- of antennomere 3. Mentum with a tooth medially, moder- belled “ Canterby N. Zeald (hand-written) / Triplosarus ately shorter than lateral lobes. Mentum and submentum fulvescens Bates (hand-written) / Ex Musaeo H.W. separated by complete transverse suture. Paraglossae as BATES 1892 (typed) / novaezelandiae cast. comparé au long as ligula. Palpi with last segment fusiform, truncate type! (hand-written) / Coll. Chaudoir (hand-written) / LECTOTYPE Triplosarus fulvescens Bates, 1878 desig- apically, with sparse, short pubescence; penultimate seg- nated by Larochelle & Larivière 2004 (red label; typed).” ment of labial palpi plurisetose on anterior margin. Tho- Perfect condition. Synonymised by Townsend, 1997: 16. rax. Pronotum transverse; base straight, moderately nar- Triplosarus novae-zealandiae [sic]: Broun, 1881: 659 (mis- rower than elytral base; lateral beads complete; anterior spelling). bead incomplete medially; posterior bead complete. Description. Body length: 7.5–10 mm. Slightly convex. Scutellum visible. Apex of prosternal lobe pubescent. Legs. Pale in colour, testaceous, sometimes moderately dark Metafemora with 5–7 long setae on posterior margin. Male brown; pronotum with light greenish tinge; antennae, protarsi and mesotarsi dilated laterally and spongily pu- mouthparts, and legs pale. Generally glabrous and smooth. bescent ventrally. Segment 4 of protarsi and mesotarsi of Microsculpture isodiametric. Pronotum shiny; head and both sexes without membranous laminae. Tarsi glabrous elytra dull; head and pronotum with bronze metallic lustre. dorsally; metatarsomere 5 pubescent (with 6 setae) Head. Big, although narrower across eyes than pronotal ventrally; metatarsomere 1 shorter than metatarsomeres apex; flat anteriorly, slightly convex posteriorly. Antennae 2+3. Elytra. Interneurs complete. Rows of setiferous punc- moderately long, reaching pronotal base; antennal scape tures absent on intervals 3, 5, and 7, and on interneur 2. about 2× longer than its maximum width. Penultimate seg- Umbilicate setiferous series of interval 9 continuous. Ab- ment of labial palpi with 4–7 long setae on anterior margin. domen. Ventrites 2+3 of male without a setiferous fovea. Thorax. Pronotum (Fig. 131) very transverse, widest be- Ventrites 5+6 of both sexes without short setae, with paired fore middle; sides converging toward base, not sinuate; ambulatory setae only. Aedeagus (Fig. 48). Lateral view: apex straight; lateral depressions widening posteriorly; strongly arcuate. Dorsal view: asymmetrical (with ostium anterior angles moderately developed, rounded; posterior strongly deflected to the right); dorsal membranous area angles strongly developed, angulate; basal foveae deep, very wide, not extending to basal bulb; apical disc present. wide; punctuation feebly developed (slightly more so in Internal sac armed. basal foveae). Apex of prosternal lobe with 4 long setae Geographic distribution. New Zealand (endemic). and 3–4 short setae. Metepisterna longer than wide. Elytra. References. Hudson, 1934: 176 (list); Noonan, 1973: 285– Widest about middle. Shoulders strongly developed, 286 and 1976: 8 (taxonomy); Larochelle & Larivière, 2001: angulate, without a tooth. Subapical sinuations feeble. 125–126 (catalogue). Sutural apices angulate. Scutellar striole present. Interneurs shallow, more or less punctate. Intervals impunctate, flat. Remarks. This monotypic genus has morphological fea- Interval 3 without setiferous puncture behind middle. tures not fitting the character complex found in New Zea- Aedeagus (Fig. 48). As for genus. land Anisodactylina: aedeagus asymmetrical with ostium strongly deflected to the right and pale body colour. It is Material examined. 173 specimens, including type also the only taxon restricted to coastal sand dune and (AMNZ, BBNZ, CMNZ, ITNZ, JNNZ, LUNZ, MCSN, beach habitats. MNHN, MONZ, NZAC, OMNZ, UCNZ). 46 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Geographic distribution (Map p. 148). North Island: ventrally (by 1.5–2× maximum width of antennal scape). AK, BP, CL, HB, ND, WI, WN, WO. South Island: DN, Tempora inflated. Frons without clypeo-ocular MC, NN, SD, SL, WD. Stewart Island. prolongations. Antennal pubescence starting from basal Ecology. Coastal lowland. Sand dunes and sand beaches. half of antennomere 3. Mentum with a tooth medially, Open ground; dry, sandy soil, bare or sparsely vegetated. moderately shorter than lateral lobes. Mentum and Crepuscular and nocturnal; sheltering during the day in submentum separated by complete transverse suture. burrows (mostly), under marram grass (Ammophila) and Paraglossae as long as ligula (clunieae, crosbyi, hallae) or wrack. Gregarious. Biology. Seasonality: October–April, longer. Palpi with last segment fusiform, truncate or not August. Tenerals: November (rarely), January–February. apically, with sparse or moderately dense long pubescence; Dispersal power. Elytra free along suture. Macropterous, penultimate segment of labial palpi plurisetose or trisetose probably capable of flight. Moderate runner. Good bur- (clunieae, hallae) on anterior margin. Thorax. Pronotum rower. Collecting techniques. Pitfall trapping, collecting transverse or suborbicular (clunieae); base emarginate, as at night with a torch, turning logs. wide as or narrower than elytral base; lateral beads com- plete; anterior bead absent; posterior bead absent or com- References. Wakefield, 1873: 298 (distribution, ecology); plete. Scutellum visible. Apex of prosternal lobe pubes- Hudson, 1934: 176 (list); Pilgrim, 1969: 364 (ecology); cent. Legs. Metafemora with 2–6 long setae on posterior Harris, 1970: 48, 55 (distribution, ecology, biology); margin. Pro-, meso-, and metatarsomeres 1–4 of both sexes Noonan, 1973: 286 (taxonomy, dispersal power); Johns, dilated laterally, and subtriangular (as opposed to 1986: 31 (distribution, ecology); Townsend, 1994: 11 (dis- Parabaris, only male pro- and mesotarsi dilated). Male tribution, ecology) and 1997: 16 (taxonomy, distribution); protarsi spongily pubescent ventrally; mesotarsi spongily Larochelle & Larivière, 2001: 125–126 (taxonomy, distri- pubescent or not (moorei) ventrally. Segment 4 of protarsi bution, ecology, biology, dispersal power). and mesotarsi of both sexes without membranous laminae. Remarks. One male specimen of Harpalus Tarsi pubescent (with numerous setae) dorsally; novaezelandiae collected from Auckland and labelled metatarsomere 5 pubescent (with numerous setae) “syntypus” was obtained from the Castelnau collection ventrally; metatarsomere 1 as long as metatarsomeres 2+3. in Genova (MCSN), it is here designated as a lectotype to Elytra. Interneurs complete. Rows of setiferous punc- preserve stability of nomenclature in the future. tures present on intervals 3, 5 or 7, or on interneur 2. Bates’ original description of Triplosarus fulvescens Umbilicate setiferous series of interval 9 continuous. Ab- was based on an unspecified number of specimens of both domen. Ventrites 2+3 of male without a setiferous fovea. sexes (“[male symbol, female symbol]”), which apparently Ventrites 5+6 of both sexes without short setae, with paired came from collectors based in “Auckland?” and ambulatory setae only. Aedeagus. Lateral view: strongly “Christchurch”. Five specimens were obtained from the arcuate. Dorsal view: asymmetrical (with ostium deflected Bates collection in Paris (MNHN), one of which (a male) slightly to the right or undulated (clunieae)); dorsal mem- was collected in “Canterby [=Canterbury] N. Zeald” branous area wide, extending almost to basal bulb; apical (Christchurch’s regional area) and bears a determination disc present or absent. Internal sac armed or unarmed. label written by Bates; this specimen is here selected as lectotype to preserve stability of nomenclature in the future. Geographic distribution. New Zealand (endemic; Three The external morphology (including body colour) of T. Kings Islands and North Island). novaezelandiae is highly variable, but the configuration of References. Britton, 1964b: 522–523, 526 (taxonomy; as the male genitalia is constant throughout its range along Parabaris (in part)); Noonan, 1976: 9 (taxonomy; as coastal areas of New Zealand. Parabaris (in part)); Larochelle & Larivière, 2001: 125 (catalogue; as Parabaris (in part)). Remarks. The generic name is derived from tui (a New Genus Tuiharpalus new genus E Zealand honey-eating bird) and Harpalus (the type genus Type species. Tuiharpalus moorei new species, by present of the tribe Harpalini). The characters unifying the species designation. of this new genus, including T. gourlayi which is trans- Description. Body length: 8.0–14.0 mm. Forebody (head ferred from Parabaris, are: forebody (head and thorax) and thorax) with sparse setiferous micropores dorsally. with sparse setiferous micropores dorsally; rows of Head. Mandibles short or moderately long, slightly or setiferous punctures on elytra; pro-, meso-, and metatarsi strongly curved forward, blunt apically. Labrum moder- of both sexes dilated laterally, subtriangular. The strongly ately transverse or strongly transverse (moorei); apex reduced, rather flat eyes together with the inflated tempora, straight or slightly emarginate medially. Eyes strongly re- suggest that representatives of this genus exhibit subterra- duced, rather flat, widely separated from buccal fissure nean behaviour. Fauna of New Zealand 53 47

Key to species of Tuiharpalus Mandibles moderatly long, slightly curved forward. La- 1 Elytral interval 7 with a row of setiferous punctures (Fig. brum moderately transverse; apex straight or slightly 99). Pronotum (Fig. 136) with lateral depressions emarginate medially. Eyes widely separated from buccal × strongly explanate throughout. [Head very large; stout fissure ventrally (by at least 2 maximum width of antennal body] ...... (p. 50)... moorei new species scape). Antennae moderately long, reaching pronotal base; antennal scape about 2× longer than its maximum width. — Elytral interval 7 without a row of setiferous punctures Paraglossae as long as ligula. Palpi broadly truncate apically, (Fig. 96–98). Pronotum (Fig. 132–135) with lateral with sparse, moderately long pubescence; penultimate seg- depressions not strongly explanate throughout. [Head ment of labial palpi plurisetose (with 4–5 long setae and at smaller; slender body] ...... 2 most 1 short seta) on anterior margin. Thorax. Pronotum 2(1) Elytral interneur 2 with a row of setiferous punctures (Fig. 132) moderately transverse, widest before middle; (Fig. 98); intervals 3 and 5 without rows of setiferous sides strongly converging toward base (more so than in punctures (Fig. 98) ...... 3 gourlayi), moderately sinuate; base emarginate, moderately — Elytral interneur 2 without a row of setiferous punctures narrower than elytral base; apex concave; lateral depres- (Fig. 96–97); intervals 3 and 5 with rows of setiferous sions not explanate; posterior bead absent; anterior angles punctures (Fig. 96–97) ...... 4 moderately developed, obtusely rounded; posterior angles strongly developed, subrectangular; basal foveae deep, wide; 3(2) Pronotum (Fig. 135) very transverse; anterior angles punctuation fine, evenly distributed. Apex of prosternal moderately developed, obtusely rounded; posterior lobe with 4 long setae and several short setae. Metepisterna angles moderately developed, obtuse. Piceous black wider than long. Legs. Metafemora with 2 long setae on ...... (p. 49)... hallae new species posterior margin. Male mesotarsi spongily pubescent — Pronotum (Fig. 134) suborbicular; anterior angles ventrally. Elytra. Widest about middle. Shoulders strongly strongly developed, acute; posterior angles feebly developed, angulate, with a tooth. Subapical sinuations developed, broadly rounded. Reddish ...... rather strong. Sutural apices angulate. Scutellar striole ab- ...... (p. 48)... clunieae new species sent. Interneurs shallow, unevenly impressed, impunctate. 4(2) Pronotum (Fig. 133) very transverse; sides slightly Intervals sparsely punctate, flat; intervals 3 and 5 with converging toward base, slightly sinuate. Body length rows of setiferous punctures. Aedeagus (Fig. 49). Lateral 12.5 mm or more ...... (p. 48)... gourlayi (Britton) view: strongly arcuate; apex broadly triangular. Dorsal view: — Pronotum (Fig. 132) less transverse; sides strongly asymmetrical (with ostium slightly deflected to the right); converging toward base, moderately sinuate. Body dorsal membranous area very wide, extending almost to length 11.0 mm or less ...(p. 47)... crosbyi new species basal bulb; apical disc present, broadly triangular; main shaft straight. Internal sac unarmed. Material examined. 7 specimens, including types (AMNZ, LUNZ, NZAC). Tuiharpalus crosbyi new species E Geographic distribution (Map p. 149). Offshore Islands: Figures 49, 96, 132, 188; Map p. 149 TH-Great Island. Tuiharpalus crosbyi Larochelle & Larivière, new species. Ecology. Lowland. Wet forests. Shaded ground. Noctur- Holotype: male (NZAC) labelled “THREE KINGS IS NZ nal; sheltering during the day under stones. Biology. Great I, 45 m 28–30 Nov. 1983 C.F. Butcher (typed) / Pan Seasonality: November–December, April. Dispersal traps Shore and coastal forest (typed) / HOLOTYPE [male power. Elytra fused along suture. Subapterous. Moderate symbol] Tuiharpalus crosbyi Larochelle & Larivière, 2004 runner. Collecting techniques. Turning stones, pitfall (red label; typed).” Paratypes: 3 males (2 NZAC, Great trapping, collecting at night with a torch, using pan traps. Island, Tasman Valley; 1 LUNZ, Great Island), bearing Remarks. This species is morphologically close to T. blue paratype labels. gourlayi with which it co-occurs on the Three Kings Is- Description. Body length: 10.5–11.0 mm. Slightly con- lands. T. crosbyi is named after our colleague Trevor K. vex. Black; pronotal margins, labrum, antennae, palpi, and Crosby (Landcare Research, Auckland) for his contribu- tarsi rufous. Microsculpture isodiametric on head, moder- tion as Editor of the Fauna of New Zealand series. ately transverse on pronotum, and very transverse (with microlines) on elytra. Moderately shiny, without metallic lustre. Head. Moderately large, narrower across eyes than pronotal apex; flat anteriorly, slightly convex posteriorly. 48 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Tuiharpalus gourlayi (Britton, 1964) new combination E Geographic distribution (Map p. 149). Offshore Islands: Figures 50, 97, 133, 189; Map p. 149 TH–Great Island. North East Island. Parabaris gourlayi Britton, 1964b: 523. Holotype: male South West Island. West Island. (NZAC) labelled “Type (circular red-bordered label; typed) / Great Island Three Kings 1–3.1.63 E. S. Gourlay (hand- Ecology. Lowland. Wet forests (broadleaf). Shaded ground. written) / HOLOTYPE [male symbol] Parabaris gourlayi Nocturnal; sheltering during the day under stones (mostly) mihi (hand-written) E. B. Britton det. 1963 (typed, ex- and logs, in leaf litter, and in nests of petrel colonies. Biol- cept for number 3).” Perfect condition. There are 10 ogy. Seasonality: November–January, May. Tenerals: No- paratypes in NZAC and there should be another 3 in vember–December (mostly), January, April. Regularly in- BMNH. fested with mites. Dispersal power. Elytra fused along Description. Body length: 12.5–14.0 mm. Slightly con- suture. Subapterous. Moderate runner. Collecting tech- vex. Black; pronotal margins dark reddish brown; antennae niques. Turning stones and logs, collecting at night with a and palpi light reddish brown. Microsculpture shallow, torch, pitfall trapping, raking leaf litter. isodiametric on head, shallow isodiametric to slightly trans- References. Britton, 1964b: 526 (distribution); Larochelle verse on pronotum, very transverse (with microlines) on & Larivière, 2001: 125 (taxonomy, distribution, ecology, elytra. Shiny (especially elytra), without metallic lustre. biology, dispersal power). Elytra slightly iridescent. Head. Moderately large, nar- rower across eyes than pronotal apex; flat anteriorly, slightly Remarks. Examination of the type of Parabaris gourlayi convex posteriorly. Mandibles short, strongly curved for- revealed it to be congeneric with taxa placed in the new ward. Labrum moderately transverse; apex slightly genus Tuiharpalus. See also Remarks under T. crosbyi. emarginate medially. Eyes widely separated from buccal fissure ventrally (by about 2× maximum width of antennal E scape). Antennae moderately long, reaching pronotal base; Tuiharpalus clunieae new species antennal scape about 2× longer than its maximum width. Figures 51, 134, 190; Map p. 149 Paraglossae longer than ligula. Palpi broadly truncate Tuiharpalus clunieae Larochelle & Larivière, new species. apically, with moderately dense and long pubescence; pe- Holotype: male (NZAC) labelled “Trounson Kauri Park, nultimate segment of labial palpi plurisetose (with 4 long Donnelly’s Crossing, ND. 25-11-94 J.I. Townsend [hand- written] / HOLOTYPE [male symbol] Tuiharpalus setae and 1 short seta) on anterior margin. Thorax. clunieae Larochelle & Larivière, 2004 (red label; typed).” Pronotum (Fig. 133) very transverse, widest before mid- Paratypes: 2 females (2 AMNZ, Waipoua Forest) bearing dle; sides slightly converging toward base (less so than in blue paratype labels. crosbyi), slightly sinuate; base emarginate, slightly nar- Description. Body length: 9.0–10.0 mm. Forebody (head rower than elytral base; apex concave; lateral depressions and thorax) strongly convex; hindbody moderately con- strongly explanate posteriorly; posterior bead absent; an- vex. Reddish (including antennae and palpi). Microsculpture terior angles moderately developed, obtusely rounded; deep, isodiametric on head, moderately transverse on posterior angles strongly developed, subrectangular; basal pronotum, and deep, isodiametric on elytra (females) or foveae deep, wide; punctuation fine, evenly distributed. granulate (males). Shiny, without metallic lustre; elytra Apex of prosternal lobe with 3–5 long setae and several dull in males. Head. Moderately large, narrower across short setae. Metepisterna wider than long. Legs. eyes than pronotal apex; flat anteriorly, slightly convex Metafemora with 4–6 long setae on posterior margin. Male posteriorly. Mandibles moderately long, slightly curved mesotarsi spongily pubescent ventrally. Elytra. Widest forward. Labrum moderately transverse; apex straight or about middle. Shoulders strongly developed, angulate, with slightly emarginate medially. Eyes widely separated from a tooth. Subapical sinuations strong. Sutural apices angulate. buccal fissure ventrally (by about 1.5× maximum width of Scutellar striole absent. Interneurs shallow, impunctate. antennal scape). Antennae moderately long, reaching about Intervals sparsely punctate, flat; intervals 3 and 5 with pronotal base; antennal scape about 2× longer than its maxi- rows of setiferous punctures. Aedeagus (Fig. 50). Lateral mum width. Paraglossae as long as ligula. Palpi not trun- view: strongly arcuate; apex narrowly triangular (more so cate apically, with sparse long pubescence; penultimate than in crosbyi). Dorsal view: asymmetrical (with ostium segment of labial palpi trisetose (with 2 long and 1 short slightly deflected to the right); dorsal membranous area seta) on anterior margin. Thorax. Pronotum (Fig. 134) very wide, extending almost to basal bulb; apical disc suborbicular, widest about middle; sides converging to- present, narrowly triangular; main shaft deflected to the ward base, not sinuate; base emarginate, moderately nar- right. Internal sac unarmed. rower than elytral base; apex concave; lateral depressions Material examined. 290 specimens, including types moderately explanate throughout; posterior bead complete; (AMNZ, BMNH, CMNZ, LUNZ, NZAC). anterior angles strongly developed, acute; posterior angles Fauna of New Zealand 53 49 feebly developed, broadly rounded; basal foveae shallow, transverse on pronotum, and deep, isodiametric on elytra wide; punctuation fine, evenly distributed. Apex of (females) or granulate on elytra (males). Shiny, without prosternal lobe with 2 long setae and several short setae. metallic lustre; elytra dull in males. Head. Moderately Metepisterna as wide as long. Legs. Metafemora with 2 large, narrower across eyes than pronotal apex; flat long setae on posterior margin. Male protarsomeres 1–4 anteriorly, slightly convex posteriorly. Mandibles moder- unusually wide, 3× wider than mesotarsomeres (contrary ately long, slightly curved forward. Labrum moderately to other Tuiharpalus species, except hallae). Male transverse; apex straight or slightly emarginate medially. mesotarsi spongily pubescent ventrally. Elytra. Widest Eyes widely separated from buccal fissure ventrally (by about middle. Shoulders well developed, rounded, with a about 1.5× maximum width of antennal scape). Antennae tooth. Subapical sinuations feeble. Sutural apices angulate. moderately long, reaching about pronotal base; antennal Scutellar striole absent or present. Interneurs shallow, scape about 2× longer than its maximum width. Paraglossae impunctate, except interneur 2 with a row of setiferous as long as ligula. Palpi narrowly truncate apically, with punctures. Intervals sparsely punctate, flat; intervals 3 sparse, moderately long pubescence; penultimate segment and 5 without rows of setiferous punctures. Aedeagus of labial palpi trisetose (with 2 long and 1 short setae) on (Fig. 51). Lateral view: strongly arcuate; apex narrowly anterior margin. Thorax. Pronotum (Fig. 135) very trans- pointed. Dorsal view: asymmetrical (with ostium undu- verse, widest about middle; sides converging toward base, lated, slightly deflected to the left about middle and to the not sinuate; base emarginate, as wide as elytral base; apex right subapically); dorsal membranous area moderately concave; lateral depressions moderately explanate through- wide, extending almost to basal bulb; apical disc present, out; posterior bead absent; anterior angles moderately de- triangular; main shaft undulated. Internal sac armed. veloped, obtusely rounded; posterior angles moderately Material examined. 5 specimens, including types developed, obtuse; basal foveae shallow, wide; punctua- (AMNZ, NZAC). tion fine, evenly distributed. Apex of prosternal lobe with 2 long setae and several sparse short setae. Metepisterna Geographic distribution (Map p. 149). North Island: as wide as long. Legs. Metafemora with 2 long setae on ND–Trounson Kauri Park. Waipoua Forest. posterior margin. Male protarsomeres 1–4 unusually wide, Ecology. Lowland. Wet forests (broadleaf, podocarp): along 3× wider than mesotarsomeres (contrary to other streams. Shaded ground; soil covered with dead leaves. Tuiharpalus species, except clunieae). Male mesotarsi Nocturnal; sheltering during the day under logs. Biology. spongily pubescent ventrally. Elytra. Widest about mid- Seasonality: October-November, January. Dispersal dle. Shoulders strongly developed, rounded, with a tooth. power. Elytra fused along suture. Subapterous. Moderate Subapical sinuations feeble. Sutural apices angulate. runner. Collecting techniques. Turning logs. Scutellar striole absent or present. Interneurs shallow, Remarks. This species is morphologically close to T. impunctate, except interneur 2 with a row of setiferous hallae. Both taxa occur between Kaitaia and the punctures. Intervals sparsely punctate, flat; intervals 3 southernmost limit of the Northland region. This taxon is and 5 without rows of setiferous punctures. Aedeagus named after our good friend and colleague Leonie H. Clunie (Fig. 52). Lateral view: strongly arcuate, apex narrowly (Landcare Research, Auckland) for her special help in our pointed (more attenuate than in clunieae). Dorsal view: entomological studies. asymmetrical (with ostium slightly deflected to the right); dorsal membranous area moderately wide, extending al- most to basal bulb; apical disc absent; main shaft mostly straight, slightly deflected to the right apically. Internal sac E Tuiharpalus hallae new species armed. Figures 52, 98, 135, 191; Map p. 149 Tuiharpalus hallae Larochelle & Larivière, new species. Material examined. 13 specimens, including types Holotype: male (NZAC) labelled “ NEW ZEALAND (AMNZ, CMNH, NZAC, OMNZ). ND Mangamuka Gorge 4.II.2004 Bob Ward / roadside Geographic distribution (Map p. 149). North Island: drainage, under leaves / HOLOTYPE [male symbol] ND–Ahipara Plateau. Mangakahia Valley. Mangamuka Tuiharpalus hallae Larochelle & Larivière, 2004 (red Gorge. Okaihau. Omahuta Forest. Parakao. Tangihua label; typed).” Paratypes: 3 females (1 CMNH, 1 NZAC, 1 OMNZ) from the same locality as the holotype, bear- Range. ing blue paratype labels. Ecology. Lowland. Wet forests (broadleaf): along mud flats, Description. Body length: 12.0–12.5 mm. Forebody (head streams, and seepages. Shaded ground; soil covered with and thorax) strongly convex, hindbody moderately con- dead leaves. Nocturnal; sheltering during the day in leaf vex. Piceous black; antennae and palpi rufous. litter and under logs. Biology. Seasonality: October–De- Microsculpture deep, isodiametric on head, moderately cember, February–March. Tenerals: September–October. 50 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Dispersal power. Elytra fused along suture. Subapterous. Elytra. Widest about middle. Shoulders moderately devel- Moderate runner. Collecting techniques. Raking leaf lit- oped, rounded, without a tooth. Subapical sinuations mod- ter, turning logs, pitfall trapping. erate. Sutural apices angulate. Scutellar striole absent. Remarks. See under T. clunieae. This species is named Interneurs moderately deep, evenly impressed, impunctate. after our good friend and colleague Grace Hall (Landcare Intervals sparsely punctate, slightly convex; intervals 3, 5, Research, Auckland) for her special help in our entomo- 7 with rows of setiferous punctures. Aedeagus (Fig. 53). logical studies and for her warm dedication to making us Lateral view: strongly arcuate; apex obtusely rounded. enjoy our life and career in New Zealand. The holotype Dorsal view: asymmetrical (with ostium slightly deflected and two of the paratypes were graciously provided by to the right); dorsal membranous area very wide, extending R.D. Ward (Tennessee, U.S.A.) and J. Nunn (Dunedin). almost to basal bulb; apical disc present, rounded-triangu- lar; main shaft straight, inflated in apical half. Internal sac unarmed. E Tuiharpalus moorei new species Material examined. 6 specimens, including types Figures 53, 99, 136, 192; Map p. 149 (AMNZ, NZAC). Tuiharpalus moorei Larochelle & Larivière, new species. Holotype: male (NZAC) labelled “NEW ZEALAND ND Geographic distribution (Map p. 149). North Island: Te Paki Res., Te Paki Trig tk, Kauri Bush 13-17. 1995 ND–Kohuronaki. Pandora. Te Paki Trig. Unuwhao. Dec. Larivière, Larochelle (typed, except month of year) Ecology. Lowland. Wet forests (broadleaf, podocarp): along / Taraire-kauri for.: streambank; dark wet litter-rich soil. streams. Shaded ground; soil covered with dead leaves. Under stones. (typed) / HOLOTYPE [male symbol] Tuiharpalus moorei Larochelle & Larivière, 2004 (red Nocturnal; sheltering during the day in leaf litter and under label; typed).” Paratypes: 1 male (MONZ), 1 female stones. Biology. Seasonality: November–December, Feb- (NZAC) from the same locality as the holotype, bearing ruary. Dispersal power. Elytra fused along suture. blue paratype labels. Subapterous. Moderate runner. Collecting techniques. Description. Body length: 8.0–8.5 mm. Strongly convex. Raking leaf litter, pitfall trapping. Piceous brown; forebody (head and thorax) light brown; Remarks. The presence of series of setiferous punctures lateral depressions of pronotum, antennae, palpi, and legs on elytral intervals 3, 5, and 7, the stout convex body, rufous. Microsculpture isodiametric and weak on head, ovate elytra, and strongly explanate lateral depressions of moderately transverse and weak on pronotum, very trans- the pronotum set this taxon apart from its congeners. This verse (with microlines) and shallow on elytra. Very shiny, is the only species of Tuiharpalus recorded from and re- without metallic lustre. Head. Very large, only slightly stricted to the tip of the Aupouri Peninsula. Tuiharpalus narrower across eyes than pronotal apex; flat anteriorly, moorei is named after our friend and colleague Barry P. slightly convex posteriorly. Mandibles moderately long, Moore (Research Associate, Australian National Collec- slightly curved forward. Labrum strongly transverse; apex tion, Canberra, Australia) for his special help and encour- straight of slightly emarginate medially. Eyes widely sepa- agement in our carabid studies. × rated from buccal fissure ventrally (by about 2 maximum . width of antennal scape). Antennae moderately long, reach- ing pronotal base; antennal scape about 2× longer than its maximum width. Paraglossae longer than ligula. Palpi not truncate apically, with moderately dense and long pubes- Subtribe HARPALINA cence; penultimate segment of labial palpi plurisetose (with Diagnosis (New Zealand). Body length: 6.0–12.0 mm. 2–3 long setae, and 1–4 short setae) on anterior margin. Frons without clypeo-ocular prolongations. Mentum with Thorax. Pronotum (Fig. 136) very transverse, widest be- a tooth medially. Mentum and submentum separated by fore middle; sides converging toward base, not sinuate; complete transverse suture. Penultimate segment of labial base emarginate, moderately narrower than elytral base; palpi plurisetose (with 4 setae or more) on anterior margin. apex concave; lateral depressions strongly explanate Apex of prosternal lobe pubescent. Male protarsi and throughout; posterior bead absent; anterior angles strongly mesotarsi dilated laterally and biseriately pubescent (with developed, subtriangular; posterior angles feebly devel- 2 rows of scale-like setae) ventrally. Metatarsomere 1 oped, broadly rounded; basal foveae absent; punctuation shorter than metatarsomeres 2+3. Umbilicate setiferous fine, evenly distributed. Apex of prosternal lobe with 3 series of interval 9 separated into two major groups with long setae and several short setae. Metepisterna wider than posterior group continuous (not divided further into two long. Legs. Metafemora with 4 long setae on posterior subgroups). Aedeagus arcuate, asymmetrical with ostium margin. Male mesotarsi not spongily pubescent ventrally. strongly deflected to the left. Fauna of New Zealand 53 51

Geographic distribution. Worldwide. References. Noonan, 1976: 31–38 (taxonomy); Larochelle References. Noonan, 1976: 28–60 (taxonomy); Moore, & Larivière, 2001: 128 (catalogue). 1977: 7–10 (key to Australian genera); Larochelle & Larivière, 2001: 128 (catalogue). Subgenus Harpalus Latreille, 1802 A Harpalus Latreille, 1802: 92 (see above). Genus Harpalus Latreille, 1802 A Amblystus Motschulsky, 1864: 209. Type species: Carabus rubripes Duftschmid, 1812, by original designation. Au- Harpalus Latreille 1802: 92. Type species: Carabus proteus thor of synonymy unknown for this subgenus. Paykull, 1790 (=Carabus affinis Schrank, 1781), desig- nated by Andrewes, 1935: 19. Geographic distribution. Same as genus. Description (New Zealand). Body length: 6.0–12.0 mm. Forebody (head and thorax) with or without sparse setiferous micropores dorsally. Head. Mandibles short or Key to species of Harpalus moderately long, strongly curved forward, blunt apically. 1 Pronotum (Fig. 137): sides converging toward base; Labrum moderately or strongly transverse; apex straight punctuation strongly developed basally and laterally; or slightly emarginate medially. Eyes moderately large, base emarginate. Elytra with apex and outer intervals convex, moderately or widely separated from buccal fis- pubescent (Fig. 100). [Metallic lustre strong; Fig. 193] sure ventrally (by 1–1.5× maximum width of antennal ...... (p. 51)... affinis (Schrank) scape). Tempora not inflated. Frons without clypeo-ocu- — Pronotum (Fig. 138–139): sides not converging toward lar prolongations. Antennal pubescence starting from mid- base; punctuation feebly developed basally and laterally; dle of antennomere 3. Mentum with a tooth medially, mod- base straight. Elytra with apex glabrous and at most erately or much shorter than lateral lobes. Mentum and interval 9 pubescent (in addition to umbilicate setiferous submentum separated by complete transverse suture. series). [Metallic lustre absent or weak] ...... 2 Paraglossae as long as or longer than ligula. Palpi with last 2(1) Body greenish dorsally (Fig. 195). Elytra with interval segment fusiform, not truncate apically, with sparse, mod- 9 glabrous (except for umbilicate series of setiferous erately long pubescence; penultimate segment of labial palpi punctures). Microsculpture strongly developed on plurisetose on anterior margin. Thorax. Pronotum very head. Paraglossae longer than ligula. Metafemora with transverse; base straight or emarginate, as wide as elytral 4 long setae on posterior margin. Body length 7.5 mm base; lateral beads complete; anterior bead incomplete me- or less. [Pronotum (Fig. 139)] ...... dially; posterior bead complete. Scutellum visible. Apex of ...... (p. 53)... australasiae Dejean prosternal lobe pubescent. Legs. Metafemora with 4–10 long setae on posterior margin. Male protarsi and mesotarsi — Body not greenish dorsally (Fig. 194). Elytra with dilated laterally and biseriately pubescent ventrally. Seg- interval 9 pubescent (in addition to umbilicate series of ment 4 of protarsi and mesotarsi of both sexes without setiferous punctures). Microsculpture almost absent membranous laminae. Tarsi glabrous (at least tarsomeres on head. Paraglossae as long as ligula (Fig. 31). 1–4) dorsally; metatarsomere 5 pubescent (with 6–8 se- Metafemora with 8-10 long setae on posterior margin. tae) ventrally; metatarsomere 1 much shorter than Body length 8.0 mm or more. [Pronotum (Fig. 138)] metatarsomeres 2+3 (slightly longer than metatarsomere ...... (p. 52)... tardus (Panzer) 2). Elytra. Interneurs complete. Rows of setiferous punc- tures absent on intervals 3, 5, and 7, and on interneur 2. A Umbilicate setiferous series of interval 9 separated into Harpalus (H.) affinis (Schrank, 1781) two major groups, with posterior group continuous. Ab- Figures 54, 100, 137, 193; Map p. 149 domen. Ventrites 2+3 of male without a setiferous fovea. Carabus aeneus Fabricius, 1775: 245. Type locality: Ger- many. Primary homonym of Carabus aeneus DeGeer, Ventrites 5+6 of both sexes without short setae, with paired 1774. ambulatory setae only. Aedeagus. Lateral view: moder- Carabus affinis Schrank, 1781: 212 (replacement name for ately arcuate. Dorsal view: asymmetrical (with ostium Carabus aeneus Fabricius, 1775). Type locality: Austria. strongly deflected to the left); dorsal membranous area Harpalus affinis: Author of combination uncertain for this very wide, extending only in apical half (stopping well European species. before basal bulb); apical disc present. Internal sac armed Description. Body length: 8.2–12.0 mm. Moderately con- or unarmed. vex. Black; antennae and legs either reddish or blackish. Geographic distribution. Nearctic, Palearctic, Ethiopian, Head and pronotum glabrous and smooth; elytra with outer Oriental, and Australian Regions; New Zealand (adventive). intervals and apex pubescent. Microsculpture isodiametric 52 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

(stronger in males), absent on frons and pronotal disc. along suture. Macropterous. Frequent flier. Moderate run- Shiny, with strong metallic lustre (often green, otherwise ner. Clearly effective as a colonist. Strongly favored by bronze or coppery); elytra duller in females. Head. Mod- human activities. Collecting techniques. Pitfall trapping, erately large, narrower across eyes than pronotal apex; flat turning stones and logs. anteriorly, slightly convex posteriorly. Mandibles moder- References. Lindroth, 1986: 351 (distribution, ecology, ately long. Labrum strongly transverse; apex straight. Eyes biology, dispersal power); Townsend, 1992: 25–29 (distri- widely separated from buccal fissure ventrally (by about bution, ecology, biology) and 1994: 9, 11 (distribution, × 1.5 maximum width of antennal scape). Antennae moder- ecology); Sunderland et al., 1995: 39–49 (biology); ately long, reaching pronotal base; antennal scape about 2× Larochelle & Larivière, 2001: 128 (taxonomy, distribution, longer than its maximum width. Mentum with medial tooth ecology, biology, dispersal power) and 2003: 273–274 (ecol- moderately shorter than lateral lobes. Paraglossae as long ogy, biology, dispersal power, collecting techniques). as ligula. Penultimate segment of labial palpi with 2 long setae and 3–4 short setae on anterior margin. Thorax. Pronotum (Fig. 137) very transverse, widest about middle; Harpalus (H.) tardus (Panzer, 1797) A sides converging toward base, not sinuate; base emarginate; Figures 55, 138, 194; Map p. 149. apex concave; lateral depressions absent; anterior angles Carabus tardus Panzer, 1797: 24. Type locality: Germany. strongly developed, broadly rounded; posterior angles Multiple synonyms exist in the Old World literature for this strongly developed, obtuse; basal foveae deep, wide; punc- adventive species. tuation strongly developed (basally and laterally). Apex of Description. Body length: 8.0–11.0 mm. Moderately prosternal lobe with 6–7 long setae and 8–9 short setae. convex. Black; antennae, palpi, and legs reddish; femora Metepisterna longer than wide. Legs. Metafemora with 5 and tarsi brownish black; sides of pronotum paler. Gla- long setae on posterior margin. Elytra. Widest behind mid- brous and smooth, except elytral interval 9 pubescent dle. Shoulders well developed, rounded, without a tooth. throughout. Microsculpture isodiametric (stronger in Subapical sinuations strong. Sutural apices obtusely males), almost absent on head; granulate on female elytra. angulate in males, acutely angulate in females. Scutellar Shiny, without metallic lustre; elytra dull in females. Head. striole present. Interneurs shallow, impunctate. Intervals Moderately large, narrower across eyes than pronotal apex; impunctate, flat; elytral apex and outer intervals sparsely flat anteriorly, slightly convex posteriorly. Mandibles mod- pubescent. Interval 3 without setiferous puncture behind erately long. Labrum strongly transverse; apex straight or middle. Aedeagus (Fig. 54). Lateral view: as for genus; slightly emarginate. Eyes widely separated from buccal apex short (more so than in tardus), harpoon-like. Dorsal fissure ventrally (by about 1.5× maximum width of antennal view: as for genus; apical disc not much longer than wide. scape). Antennae moderately long, reaching pronotal base; Internal sac unarmed. antennal scape about 2× longer than its maximum width. Mentum with medial tooth much shorter than lateral lobes. Material examined. 70 non-type specimens (AMNZ, Paraglossae as long as ligula. Penultimate segment of labial ITNZ, JNNZ, MONZ, NZAC, OMNZ, UCNZ). palpi with 2 long setae and 3–4 short setae on anterior Geographic distribution (Map p. 149). North Island: margin. Thorax. Pronotum (Fig. 138) very transverse, RI, WA, WI, WN. Adventive. Extralimital range: Europe, widest in basal half; sides not converging toward base, not Asia; North America. First New Zealand record: Palmerston sinuate; base straight; apex concave; lateral depressions North, WI, 1975 (NZAC; Townsend, 1992: 25). Well es- widening posteriorly; anterior angles strongly developed, tablished. broadly rounded; posterior angles strongly developed, Ecology. Lowland, montane, subalpine, alpine. Vicinities subrectangular; basal foveae shallow, narrow; punctuation of cities. Cultivated fields (seradella, pea, wheat, quinoa, feebly developed (in basal foveae and around posterior buckwheat, phacelia, coriander, barley), pastures, mead- angles). Apex of prosternal lobe with 6–7 long setae and 8– ows, roadsides, gardens, sand dunes, kiwifruit orchards; 9 short setae. Metepisterna longer than wide. Legs. glasshouses and houses (occasionally). Open ground; sandy Metafemora with 8–10 long setae on posterior margin. (mostly), silty or clayey soil, covered with grass or weeds. Elytra. Widest behind middle. Shoulders well developed, Mostly nocturnal; sheltering during the day in the soil at angulate, with a strong tooth. Subapical sinuations feeble. the base of plants and under stones. Biology. Seasonality: Sutural apices obtusely angulate in males, acutely angulate throughout the year, except May–June. Spring breeder. in females. Scutellar striole present. Interneurs shallow, Tenerals: February–May. Adult overwinterer. Omnivorous, impunctate. Intervals impunctate, flat; interval 9 pubes- mostly phytophagous. Adult food, in the field: weed seeds cent (in addition to umbilicate setiferous series). Interval 3 (mostly), aphids, and flies. Dispersal power. Elytra free with a setiferous puncture behind middle. Aedeagus (Fig. Fauna of New Zealand 53 53

55). Lateral view: as for genus; apex long (more so than in longer than ligula. Penultimate segment of labial palpi with affinis), harpoon-like (in lateral view). Dorsal view: as for 2–3 long setae and 1–5 short setae on anterior margin. genus; apical disc much longer than wide. Internal sac un- Thorax. Pronotum (Fig. 139) very transverse, widest in armed. basal 1/2; sides not converging toward base, not sinuate; Material examined. 10 non-type specimens (LUNZ, base straight; apex concave; lateral depressions absent; NZAC). anterior angles strongly developed, broadly rounded; pos- terior angles strongly developed, subrectangular; basal Geographic distribution (Map. p. 149). South Island: foveae shallow, narrow; punctuation feebly developed (al- MC–Christchurch, Mount Pleasant. Adventive. most absent). Apex of prosternal lobe with 3–8 long setae Extralimital distribution: Palearctic Region. First New Zea- and 6–9 short setae. Metepisterna longer than wide. Legs. land record: 1995 (Emberson, 2004; LUNZ). Metafemora with 4 long setae on posterior margin. Elytra. Ecology. Lowland. An open rocky spur with loamy soil Widest about middle. Shoulders strongly developed, covered with grass and herbs. Nocturnal and diurnal; active angulate, with a tooth. Subapical sinuations feeble. Sutural during the day on bare ground, on pavement, and on apices angulate. Scutellar striole present. Interneurs shal- roadsides, or sheltering under stones and in burrows. Eu- low, deepening apically, impunctate. Intervals impunctate, rope: dunes, grasslands, heaths, and cultivated fields; open flat, becoming convex apically; interval 9 glabrous (except ground, with rather dry, sandy soil. Biology. Seasonality: for umbilicate setiferous series). Interval 3 with a setiferous September–October, December–January, March. Spring puncture behind middle. Aedeagus (Fig. 56). Lateral view: breeder and adult overwinterer (Europe). Dispersal power. as for genus; apex slender, not harpoon-like. Dorsal view: Elytra free along suture. Macropterous, capable of flight. apical disc moderately longer (about 1.5×) than wide. In- Moderate runner. Collecting techniques. Pitfall trapping. ternal sac armed. References. Lindroth, 1986: 362 (ecology, biology); Material examined. 198 non-type specimens (AMNZ, Emberson, 2004 (distribution, ecology, biology, dispersal ITNZ, JNNZ, LUNZ, MONZ, NZAC, OMNZ). power). Geographic distribution (Map p. 149). North Island: BP, GB, HB, WA, WI, WN. South Island: MC Subgenus (Uncertain) (Christchurch, 1960). Extralimital range: Australia (includ- ing Tasmania). Adventive. First New Zealand records: New Harpalus australasiae Dejean, 1829 reinstatement A Zealand (Redtenbacher, 1868: 15); Napier, HB 1947 Figures 56, 101, 139, 195; Map p. 149 (NZAC). Well established. Harpalus australasiae Dejean, 1829: 386. Type locality: Ecology. Lowland. Cultivated fields (strawberry), vacant Australia (as Nouvelle-Hollande). lots, lawns, and sand dunes. Open ground; dry soil covered Hypharpax australasiae: Bates, 1874: 272. Diaphoromerus australasiae: Chaudoir, 1878: 480. with sparse vegetation (grass, weeds, herbs). Mostly diur- Notiobia (Anisotarsus) australasiae: Noonan, 1973: 296. nal; active on plants in the sunshine; sheltering on cloudy Hypharpax australasiae: Moore, in Moore et al., 1987: 237; days in burrows dug at the base of plants (e.g., Lupinus, Larochelle & Larivière, 2001: 124. Muhlenbeckia) and under logs. Gregarious. Biology. Description. Body length: 6.0–7.5 mm. Slightly convex. Seasonality: throughout the year, except May–June. Black; pronotum and elytra greenish; pronotal margins red- Tenerals: January–February. Omnivorous, probably dish; base of antennae and tibiae testaceous; remainder of granivorous (Moore et al., 1987: 237). Once recorded feed- legs and palpi dark brown. Generally glabrous and smooth. ing on strawberry seeds. Dispersal power. Elytra free Microsculpture deep, strongly isodiametric on head, mod- along suture. Macropterous. Occasional flier. Moderate erately isodiametric on pronotum, slightly transverse on runner. Clearly effective as a colonist. Favoured by human elytra. Shiny, with bronze metallic lustre (males); duller activities. Collecting techniques. Pitfall trapping, dig- (females). Head. Moderately large, narrower across eyes ging at the base of plants, raking leaf litter, turning seashore than pronotal apex; flat anteriorly, slightly convex debris. posteriorly. Mandibles short. Labrum moderately trans- References. Thomson, 1922: 284 (distribution); Pilgrim, verse; apex straight or slightly emarginate medially. Eyes 1963: 841 (distribution); Noonan, 1973: 296 and 1976: 10 moderately separated from buccal fissure ventrally (by (taxonomy); Moore et al., 1987: 237 (distribution, ecol- about maximum width of antennal scape). Antennae mod- ogy, biology, dispersal power); Townsend, 1994: 9, 11, 13 erately long, reaching pronotal base; antennal scape about (taxonomy, distribution, ecology); Larochelle & Larivière, 2× longer than its maximum width. Mentum with medial 2001: 124 (as Hypharpax; taxonomy, distribution, ecol- tooth moderately shorter than lateral lobes. Paraglossae ogy, biology, dispersal power). 54 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Remarks. This species has been previously placed in — Apex of prosternal lobe glabrous (Fig. 2). Penultimate Anisodactylina, but the presence of biseriately pubescent segment of labial palpi bisetose on anterior margin (Fig. male tarsi and the aedeagal ostium strongly deflected to the 11). Eyes reaching buccal fissure (Fig. 21) or narrowly left indicate that it belongs to the Harpalina. The species separated from it ventrally (by 0.3–1× maximum width agrees in other morphological characters with members of of antennal scape; Fig. 20) ...... 2 the genus Harpalus. 2(1) Segment 4 of protarsi and mesotarsi with 2 membranous laminae (Fig. 25). Forebody (head and thorax) much narrower than elytra (Fig. 211–213) ...... (p. 68)... Syllectus Bates Subtribe PELMATELLINA — Segment 4 of protarsi and mesotarsi without Diagnosis (New Zealand). Body length: 3.2–10.0 mm. membranous laminae (Fig. 26). Forebody (head and Frons usually with clypeo-ocular prolongations, seldom thorax) at most moderately narrower than elytra (Fig. without (Syllectus gouleti). Mentum with a tooth medially. 196–199, 203–210) ...... 3 Mentum and submentum separated by complete trans- 3(2) Eyes strongly reduced (Fig. 102). Mandibles very verse suture. Penultimate segment of labial palpi usually long (about 5× their maximum width; Fig. 102). Elytral bisetose (with 2 setae), seldom trisetose (with 3 setae, interneurs absent or incomplete basally (Fig. 102). Kupeharpalus) on anterior margin. Apex of prosternal lobe Pronotum cordate or subcordate (Fig. 140–143) [South usually glabrous, seldom pubescent (Kupeharpalus). Male Island: NN, SD] ...(p. 54)... Hakaharpalus new genus protarsi dilated laterally and spongily pubescent ventrally; — Eyes normally developed (Fig. 107). Mandibles shorter male mesotarsi usually dilated laterally, spongily pubes- (Fig. 107). Elytral interneurs complete (Fig. 107). cent ventrally (except Syllectus), seldom unmodified (nei- ther dilated nor spongily pubescent; Kupeharpalus johnsi, Pronotum neither cordate nor subcordate (Fig. 147– 154). [Throughout New Zealand] ...... Lecanomerus marrisi). Metatarsomere 1 usually as long ...... (p. 60)... Lecanomerus Chaudoir as metatarsomeres 2+3, rarely shorter (Lecanomerus atriceps, L. latimanus, L. marrisi). Umbilicate setiferous series of interval 9 separated into 2 major groups with Genus Hakaharpalus new genus E posterior group either divided further into 2 subgroups or Type species. Hakaharpalus rhodeae new species, by present continuous (Hakaharpalus, Kupeharpalus, Lecanomerus designation. insignitus). Aedeagus usually arcuate, seldom almost straight Description. Body length: 3.7–4.9 mm. Forebody (head (some Syllectus), symmetrical (with ostium dorsal, not and thorax) without sparse setiferous micropores dorsally. deflected laterally). Head. Dorsal surface excavated anteriorly (as in Geographic distribution. Mostly Neotropical and Aus- Lecanomerus marrisi). Mandibles very long (about 5× tralian Regions; also Nearctic Region. their maximum width), slightly curved forward, acute References. Noonan, 1976: 6–8 (taxonomy); Larochelle apically. Labrum strongly transverse; apex slightly & Larivière, 2001: 118–122 (catalogue). emarginate medially. Eyes strongly reduced, flat or slightly convex, consisting of obliterated facets, narrowly sepa- Remarks. All world genera recognised so far within the rated from buccal fissure ventrally (by 0.7–1× maximum Pelmatellina have been characterised by the glabrous apex width of antennal scape). Tempora not inflated. Frons with of the prosternal lobe. Kupeharpalus (new genus including clypeo-ocular prolongations incomplete toward eyes. An- 3 species) which is apparently very close to Lecanomerus, tennae widening from base to apex (contrary to other deviates from this taxonomic concept by having the apex Pelmatellina genera); pubescence starting from antennomere of the prosternal lobe pubescent. 2. Mentum with a tooth medially, as long as lateral lobes. Mentum and submentum separated by complete trans- verse suture. Paraglossae longer than ligula. Palpi hirsute Key to genera of New Zealand Pelmatellina (contrary to other Pelmatellina genera), with last segment very inflated, not truncate but needle-shaped apically, with 1 Apex of prosternal lobe pubescent. Penultimate segment very dense, moderately long pubescence; penultimate seg- of labial palpi trisetose on anterior margin (Fig. 10). ment of labial palpi bisetose on anterior margin. Thorax. Eyes widely separated from buccal fissure ventrally Pronotum subcordate or cordate; base straight, much nar- × (by 1.5–2 maximum width of antennal scape; Fig. 19) rower than elytral base; lateral beads complete; anterior [North Island: Northland] ...... and posterior beads incomplete medially. Scutellum vis- ...... (p. 57)... Kupeharpalus new genus ible. Apex of prosternal lobe glabrous. Legs. Metafemora Fauna of New Zealand 53 55 with 5 long setae on posterior margin. Male protarsi and sinuate laterally in front of posterior angle. [Aedeagus mesotarsi dilated laterally and spongily pubescent ventrally. (Fig. 57): apical half slightly curved (in lateral view); Segment 4 of protarsi and mesotarsi of both sexes without apex broadly rounded (in dorsal view)] ...... membranous laminae. Tarsi pubescent (with numerous se- ...... (p. 55)... patricki new species tae) dorsally; metatarsomere 5 pubescent (with 4 setae) 4(1) Elytral interneurs absent (Fig. 199). Pronotum (Fig. ventrally; metatarsomere 1 as long as metatarsomeres 2+3. 143) cordate, slightly sinuate laterally in front of Elytra. Interneurs absent or incomplete basally. Rows of posterior angles; anterior angles feebly developed, setiferous punctures absent on intervals 3, 5, and 7, and on rounded. Microsculpture absent on pronotum. Eyes interneur 2. Umbilicate setiferous series of interval 9 sepa- rated into two major groups, with posterior group con- flat. [Aedeagus (Fig. 60)] ...... tinuous. Abdomen. Ventrites 2+3 of male without a ...... (p. 57)... rhodeae new species setiferous fovea. Ventrites 5+6 of both sexes without short —Elytral interneurs present, although weakly impressed setae, with paired ambulatory setae only. Aedeagus. Lat- (Fig. 198). Pronotum (Fig. 142) subcordate, not sinuate eral view: strongly arcuate. Dorsal view: symmetrical (with laterally in front of posterior angles; anterior angles ostium dorsal, not deflected laterally); dorsal membranous strongly developed, acute. Microsculpture present on area wide, extending to basal bulb; apical disc absent. Inter- pronotum. Eyes slightly convex. [Aedeagus (Fig. 59)] nal sac unarmed...... (p. 56)... davidsoni new species Geographic distribution. New Zealand (endemic; South Island). Remarks. The generic name is derived from haka (tradi- tional Maori chant of defiance accompanied by stylised Hakaharpalus patricki new species E movements of hands and feet) and Harpalus (the type Figures 57, 102, 140, 196; Map p. 150 genus of the tribe Harpalini). A very distinctive genus re- Hakaharpalus patricki Larochelle & Larivière, new species. stricted to the BR–NN–SD regions, which is easily recog- Holotype: male (NZAC) labelled “Mt Domett NN. 1250m nised by the more or less heart-shaped pronotum, ovate Nov-Dec 71 G. Kuschel (typed) / moss (typed) / elytra, palpi hirsute, with the last segment very inflated HOLOTYPE [male symbol] Hakaharpalus patricki and needle-shaped apically, strongly reduced eyes, and Larochelle & Larivière, 2004 (red label; typed).” antennae widening from base to apex. The strongly re- Paratypes: 2 males (1 OMNZ, 1 NZAC) and 1 female (NZAC) from the same locality as the holotype, bearing duced eyes, long pubescence, and slightly convex body blue paratype labels. suggest subterranean behaviour similar to that of Anillina (Bembidiini) that live deep in thick leaf litter and/or in soil Description. Body length: 4.1–4.5 mm. Slightly convex. fissures. Blackish brown; margins and sutures of elytra, as well as antennae, palpi, and legs pale yellowish. Generally gla- brous. Elytral intervals coarsely punctate; interneurs shal- low (weakly impressed), incomplete basally. Key to species of Hakaharpalus Microsculpture absent on head, moderately transverse on 1 Elytral intervals punctate (Fig. 102)...... ….. 2 pronotum, and very transverse (with microlines) on elytra. —Elytral intervals impunctate...... ….. 4 Shiny, without metallic lustre. Head. Moderately large, narrower across eyes than pronotal apex; excavated 2(1) Elytra (Fig. 197): interneurs deep (strongly impressed). anteriorly, slightly convex posteriorly. Eyes strongly re- [Intervals finely punctate. Aedeagus (Fig. 58): apical duced, slightly convex, consisting of obliterated facets, half almost straight (in lateral view); apex narrowly narrowly separated from buccal fissure ventrally (by about rounded (in dorsal view). Pronotum (Fig. 141)]...... 0.7× maximum width of antennal scape). Antennal scape ...... (p. 56)... maddisoni new species about 3× longer than its maximum width. Thorax. Pronotum —Elytra (Fig. 102, 196): interneurs shallow (weakly (Fig. 140) subcordate, widest before middle; sides con- impressed) ...... 3 verging toward base, not sinuate; base straight; apex al- 3(2) Microsculpture absent on pronotum and elytra. Elytral most straight behind anterior angles; lateral depressions intervals finely punctate. Pronotum (Fig. Addendum) absent; anterior angles strongly developed, acute; poste- sinuate laterally in front of posterior angle. [Male rior angles moderately developed, obtuse; basal foveae shal- low, ill-defined; punctuation feebly developed. unknown] ...... Metepisterna wider than long. Elytra. Widest about mid- ...... (p. 57, 93)...cavelli (Broun) new combination dle. Shoulders feebly developed, rounded, without a tooth. —Microsculpture present on pronotum and elytra. Elytral Subapical sinuations feeble. Sutural apices rounded. intervals coarsely punctate. Pronotum (Fig. 140) not Scutellar striole absent. Interneurs shallow, impunctate, 56 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) incomplete basally. Intervals coarsely punctate, flat. Inter- Subapical sinuations feeble. Sutural apices rounded. val 3 without setiferous puncture behind middle. Aedeagus Scutellar striole absent. Interneurs deep, impunctate, in- (Fig. 57). Lateral view: as for genus; apical half slightly complete basally. Intervals finely punctate, flat. Interval 3 curved. Dorsal view: as for genus; apex broadly rounded. without setiferous puncture behind middle. Aedeagus (Fig. Material examined. 7 specimens, including types (ITNZ, 58). Lateral view: as for genus; apical half almost straight. NZAC). Dorsal view: as for genus; apex narrowly rounded. Geographic distribution (Map p. 150). South Island: Material examined. 4 specimens, including types NN–Brown Cow (Boulder Lake Track). Calphurnia Peak (AMNZ, JNNZ, NZAC). (near Boulder Lake). Mount Domett. Geographic distribution (Map p. 150). South Island: Ecology. Montane. Wet forests (beech). Shaded ground. NN–Glasgow Range. Karamea Bluff. Nocturnal; sheltering during the day in leaf litter and moss. Ecology. Montane. Wet forests (beech). Shaded ground. Biology. Seasonality: November–January. Dispersal Biology. Seasonality: October, January–February. Dis- power. Elytra fused along suture. Subapterous. Moderate persal power. Elytra fused along suture. Subapterous. runner. Collecting techniques. Sifting leaf litter and moss. Moderate runner. Remarks. This taxon is morphologically close to H. Remarks. This species is named after our good friend maddisoni. It is named after our good colleague Brian H. Peter A. Maddison (Field Studies, Waitakere City) for his Patrick (Otago Museum, Dunedin) for his encouragement special help and encouragement in establishing our new life in our New Zealand insect studies. and career in New Zealand. J. Nunn (Dunedin) graciously provided the female paratype. See also Remarks under H. patricki. Hakaharpalus maddisoni new species E Figures 58, 141, 197; Map p. 150 Hakaharpalus maddisoni Larochelle & Larivière, new spe- Hakaharpalus davidsoni new species E cies. Holotype: male (NZAC) labelled “Surveyers Ck. Figures 59, 142, 198; Map p. 149 380m. Karamea Bluff Nelson. (hand-written) / 13 Oct Hakaharpalus davidsoni Larochelle & Larivière, new spe- 70 JI Townsend (hand-written) / HOLOTYPE [male cies. Holotype: male (NZAC) labelled “Gowan Bridge symbol] Hakaharpalus maddisoni Larochelle & Buller gorge Wet moss 24.9.64 J.I. Townsend (hand-writ- Larivière, 2004 (red label; typed).” Paratypes: 1 male ten) / HOLOTYPE [male symbol] Hakaharpalus from same locality as holotype (NZAC) and 1 female davidsoni Larochelle & Larivière, 2004 (red label; (OMNZ) from Karamea Bluff. typed).” Paratype: 1 female (NZAC) from Mt Arthur, Description. Body length: 4.2–4.5 mm. Slightly convex. NN, bearing blue paratype label. Blackish brown; margins and sutures of elytra, as well as Description. Body length: 3.7–4.9 mm. Slightly convex. antennae, palpi, and legs pale yellowish. Generally gla- Reddish or blackish brown; disc of head, pronotum, and brous. Elytral intervals finely punctate; interneurs deep, elytra darker; antennae, palpi, and legs pale yellowish. incomplete basally. Microsculpture absent on head, mod- Generally glabrous. Elytral intervals impunctate; interneurs erately transverse on pronotum, and very transverse (with shallow (weakly impressed), incomplete basally. microlines) on elytra. Head shiny; pronotum and elytra Microsculpture absent on head and elytra; shallow and dull; dorsal surface without metallic lustre. Head. Moder- very transverse (with microlines) on pronotum. Shiny, ately large, narrower across eyes than pronotal apex; exca- without metallic lustre. Head. Moderately large, narrower vated anteriorly, slightly convex posteriorly. Eyes strongly across eyes than pronotal apex; excavated anteriorly, reduced, slightly convex, consisting of obliterated facets, slightly convex posteriorly. Eyes strongly reduced, slightly narrowly separated from buccal fissure ventrally (by about convex, consisting of obliterated facets, moderately sepa- 0.7× maximum width of antennal scape). Antennal scape rated from buccal fissure ventrally (by about maximum about 3× longer than its maximum width. Thorax. Pronotum width of antennal scape). Antennal scape about 2× longer (Fig. 141) subcordate, widest before middle; sides con- than its maximum width. Thorax. Pronotum (Fig. 142) verging toward base, not sinuate; base straight; apex al- subcordate, widest before middle; sides converging toward most straight behind anterior angles; lateral depressions base, not sinuate; base straight; apex almost straight be- absent; anterior angles strongly developed, acute; poste- hind anterior angles; lateral depressions absent; anterior rior angles moderately developed, obtuse; basal foveae shal- angles strongly developed, acute; posterior angles moder- low, ill-defined; punctuation feebly developed. ately developed, obtuse; basal foveae shallow, ill-defined; Metepisterna wider than long. Elytra. Widest about mid- punctuation feebly developed. Metepisterna wider than dle. Shoulders feebly developed, rounded, without a tooth. long. Elytra. Widest about middle. Shoulders feebly devel- Fauna of New Zealand 53 57 oped, rounded, without a tooth. Subapical sinuations fee- ventrally (by about maximum width of antennal scape). ble. Sutural apices rounded. Scutellar striole absent. Antennal scape about 2× longer than its maximum width. Interneurs shallow (weakly impressed), impunctate, in- Thorax. Pronotum (Fig. 143) cordate, widest before mid- complete basally. Intervals impunctate, flat. Interval 3 with- dle; sides converging toward base, slightly sinuate; base out setiferous puncture behind middle. Aedeagus (Fig. straight; apex slightly convex; lateral depressions absent; 59). Lateral view: as for genus; apical half strongly curved, anterior angles feebly developed, rounded; posterior an- wider than in patricki and maddisoni. Dorsal view: as for gles moderately developed, obtuse; basal foveae shallow, genus; apex moderately rounded. ill-defined; punctuation feebly developed. Metepisterna Material examined. 2 type specimens (NZAC). wider than long. Elytra. Widest about middle. Shoulders feebly developed, rounded, without a tooth. Subapical Geographic distribution (Map p. 149). South Island: sinuations feeble. Sutural apices angulate. Scutellar striole NN–Buller Gorge, Gowanbridge. Mount Arthur. absent. Interneurs absent. Intervals impunctate, flat. Inter- Ecology. Lowland, montane. A forest (beech). Biology. val 3 without setiferous puncture behind middle. Aedeagus Seasonality: September. Dispersal power. Elytra fused (Fig. 60). Lateral view: as for genus; apical half straight. along suture. Subapterous. Moderate runner. Dorsal view: as for genus; apex triangular. Remarks. This species is morphologically closer to H. Material examined. 11 specimens, including types (ITNZ, patricki and H. maddisoni than to H. rhodeae. It is named NZAC). after Robert L. Davidson (Carnegie Museum, Pittsburgh, Geographic distribution (Map p. 150). South Island: Pennsylvania), long time friend and colleague of the first NN–Maitai Valley. Upper Maitai [Valley]. SD–Port Ligar. author, for special help and encouragement in his carabid Titirangi [Bay]. research. Ecology. Lowland. Wet forests (broadleaf). Shaded ground. Nocturnal; sheltering during the day in leaf litter. Biology. Hakaharpalus cavelli (Broun, 1893) new combination E Seasonality: October, March–May. Tenerals: October, March. Dispersal power. Elytra fused along suture. See Addendum, page 93. Subapterous. Moderate runner. Collecting techniques. Note. The holotype of the species originally described as Sifting leaf litter. Tachys cavelli was examined in early April 2005 in the course of another study. The tribal and generic placement of the Remarks. The absence of dorsal body microsculpture, the species was then discovered. Because this Fauna N.Z. contri- heart-shaped pronotum, and the configuration of the bution was in an advanced stage of final production the H. aedeagus set this species apart from its congeners. The cavelli description and notes are in the Addendum. species is named after our close friend and colleague Birgit Rhode (Landcare Research, Auckland) for her special help and encouragement in our carabid studies, and for her ex- E Hakaharpalus rhodeae new species ceptional dedication as a research assistant to the second Figures 60, 143, 199; Map p. 150 author. Hakaharpalus rhodeae Larochelle & Larivière, new species. Holotype: male (NZAC) labelled “Maitai Valley Nelson 25.5.90 J.I. Townsend (hand-written) / JI Townsend Col- Genus Kupeharpalus new genus E lection (typed) / HOLOTYPE [male symbol] Hakaharpalus rhodeae Larochelle & Larivière, 2004 Type species. Kupeharpalus barrattae new species, by present (red label; typed).” Paratypes: 2 females (1MONZ, 1 designation. NZAC) from the same locality as the holotype, bearing Description. Body length: 5.0–8.5 mm. Forebody (head blue paratype labels. and thorax) without sparse setiferous micropores dorsally. Description. Body length: 4.0–4.5 mm. Slightly convex. Head. Mandibles moderately long, slightly curved for- Reddish brown; disc of head, pronotum, and elytra darker; ward, acute apically. Labrum strongly or moderately trans- antennae, palpi, and legs pale yellowish. Generally gla- verse; apex straight or slightly emarginate medially. Eyes brous. Elytral intervals impunctate; inteneurs absent. moderately large, convex, widely separated from buccal Microsculpture absent on head and pronotum, very trans- fissure ventrally (by 1.5–2× maximum width of antennal verse (with microlines) on elytra. Shiny, without metallic scape). Tempora not inflated. Frons with clypeo-ocular lustre. Head. Moderately large, narrower across eyes than prolongations complete or incomplete toward eyes. pronotal apex; excavated anteriorly, slightly convex Antennal pubescence starting on basal 1/2 of antennomere posteriorly. Eyes strongly reduced, flat, consisting of ob- 3. Mentum with a tooth medially, moderately shorter than literated facets, moderately separated from buccal fissure lateral lobes. Mentum and submentum separated by com- 58 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) plete transverse suture. Paraglossae longer than or as long 2(1) Pronotum (Fig. 145): sides strongly convex. Body as ligula (johnsi). Palpi with last segment fusiform, not length 6.0–6.5 mm. Aedeagus (Fig. 62): apex triangular truncate apically, sparsely pubescent (with moderately [Northland: tip of Aupouri Peninsula] ...... long setae); penultimate segment of labial palpi trisetose ...... (p. 59)... embersoni new species on anterior margin. Thorax. Pronotum transverse; base — Pronotum (Fig. 144): sides moderately convex. Body straight or emarginate, moderately narrower than or as wide length 7.0–8.5 mm. Aedeagus (Fig. 61): apex finger- as elytral base; lateral beads complete; anterior bead com- like [Northland: south of Aupouri Peninsula] ...... plete or incomplete medially; posterior bead incomplete ...... (p. 58)... barrattae new species medially or complete. Scutellum visible. Apex of prosternal lobe pubescent. Legs. Metafemora with 2 long setae on posterior margin. Male protarsi dilated laterally and Kupeharpalus barrattae new species E spongily pubescent ventrally. Male mesotarsi dilated lat- Figures 61, 103, 144, 200; Map p. 150. erally and spongily pubescent ventrally (with spongy pu- Kupeharpalus barrattae Larochelle & Larivière, new spe- bescence not uniformly distributed, contrary to cies. Holotype: male (NZAC) labelled “NEW ZEALAND Lecanomerus) or unmodified (johnsi). Segment 4 of protarsi ND Mangamuka Gorge Wlkwy 625m 351228S 1732640E 17.IX.-16.X.1999 Larivière, Larochelle (typed) / Wet and mesotarsi of both sexes without membranous laminae. broadleaf forest. Pittraps. (typed) / HOLOTYPE [male Tarsi pubescent (with numerous setae) dorsally; symbol] Kupeharpalus barrattae Larochelle & Larivière, metatarsomere 5 pubescent (with 5–8 setae) ventrally; 2004 (red label; typed).” Paratypes: 1 male (MONZ) metatarsomere 1 as long as metatarsomeres 2+3. Elytra. from the same locality as the holotype, 3 females (1 Interneurs complete. Rows of setiferous punctures absent MONZ, 2 NZAC) from Puketi Forest, ND, bearing blue on intervals 3, 5, and 7, and on interneur 2. Umbilicate paratype labels. setiferous series of interval 9 separated into two major Description. Body length: 7.0–8.5 mm. Strongly convex. groups, with posterior group continuous. Abdomen. Piceous brown; antennae, palpi, and legs yellowish. Gen- Ventrites 2+3 of male without a setiferous fovea. Ventrites erally glabrous and smooth. Microsculpture isodiametric 5+6 of both sexes without short setae, with paired ambu- on head, moderately transverse on pronotum, very trans- latory setae only. Aedeagus. Lateral view: moderately or verse (with microlines) on elytra. Head and pronotum strongly arcuate. Dorsal view: symmetrical (with ostium moderately shiny; elytra less shiny, iridescent; without dorsal, not deflected laterally); dorsal membranous area metallic lustre. Head. Moderately large, narrower across very wide, extending almost to basal bulb; apical disc present eyes than pronotal apex; flat anteriorly, slightly convex or absent. Internal sac armed or unarmed. posteriorly. Labrum strongly transverse; apex slightly Geographic distribution. New Zealand (endemic). emarginate. Eyes widely separated from buccal fissure ventrally (by about 1.5× maximum width of antennal scape). Remarks. The generic name is derived from Kupe (the Frons with clypeo-ocular prolongations incomplete to- legendary Polynesian navigator to whom is attributed the ward eyes. Antennae moderately long, reaching pronotal discovery of New Zealand) and Harpalus (the type genus base; antennal scape about 2× longer than its maximum of the tribe Harpalini). Members of this new genus super- width. Paraglossae longer than ligula. Penultimate segment ficially resemble those of Lecanomerus, but can be sepa- of labial palpi with 2 long setae and 1 short seta on anterior rated from the latter by the following characters: eyes widely margin. Thorax. Pronotum (Fig. 144) very transverse, separated from buccal fissure; penultimate segment of la- widest about middle; sides converging toward base, not bial palpi trisetose; apex of proternal lobe pubescent. sinuate, moderately convex; base emarginate, as wide as elytral base; apex slightly concave; lateral depressions wid- Key to species of Kupeharpalus ening posteriorly; anterior bead complete; posterior bead incomplete medially; anterior angles moderately developed, 1 Pronotum (Fig. 146): base straight; posterior angles rounded; posterior angles strongly developed, broadly subrectangular; anterior angles obtuse; punctuation rounded; basal foveae shallow, narrow; punctuation feebly strongly developed basally. Paraglossae as long as ligula developed. Apex of prosternal lobe with 10–20 short setae (Fig. 31) [Body length 6.0 mm or less. Aedeagus (Fig. (without long setae). Metepisterna slightly wider than long, 63)] ...... (p. 59)... johnsi new species almost square. Legs. Male mesotarsi dilated laterally and — Pronotum (Fig. 144–145): base emarginate; posterior spongily pubescent ventrally. Metatarsomere 5 pubescent angles broadly rounded; anterior angles rounded; (with 8 setae) ventrally. Elytra. Widest about middle. Shoul- punctuation feebly developed basally. Paraglossae ders strongly developed, rounded, without a tooth. Sub- longer than ligula [Body length 6.0 mm or more] ... 2 apical sinuations feeble. Sutural apices angulate. Scutellar Fauna of New Zealand 53 59 striole absent. Interneurs moderately deep, impunctate. than ligula. Penultimate segment of labial palpi with 2 long Intervals impunctate, slightly convex. Interval 3 without setae and 1 short seta on anterior margin. Thorax. Pronotum setiferous puncture behind middle. Aedeagus (Fig. 61). (Fig. 145) very transverse, widest about middle; sides con- Lateral view: as for genus; moderately arcuate; apex verging toward base, not sinuate, strongly convex; base abruptly narrowed into finger-like process. Dorsal view: emarginate, as wide as elytral base; apex slightly concave; as for genus; apical disc present, narrow, parallel-sided. lateral depressions widening posteriorly; anterior bead com- Material examined. 22 specimens, including types plete; posterior bead incomplete medially; anterior angles (AMNZ, FMNH, MONZ, NZAC). moderately developed, rounded; posterior angles strongly developed, broadly rounded; basal foveae shallow, narrow; Geographic distribution (Map p. 150). North Island: punctuation feebly developed. Apex of prosternal lobe AK, CL, ND. with 10–20 short setae (without long setae). Metepisterna Ecology. Lowland. Wet forests (broadleaf, podocarp). slightly wider than long, almost square. Legs. Male Shaded ground; soil covered with dead leaves. Nocturnal; mesotarsi dilated laterally and spongily pubescent ventrally. sheltering during the day in leaf litter, under logs and stones. Metatarsomere 5 pubescent (with 8 setae) ventrally. Elytra. Biology. Seasonality: September–February, May–August. Widest about middle. Shoulders strongly developed, Tenerals: October, May. Dispersal power. Elytra fused rounded, without a tooth. Subapical sinuations feeble. along suture. Subapterous. Moderate runner. Collecting Sutural apices angulate. Scutellar striole absent. Interneurs techniques. Pitfall trapping, raking leaf litter, turning logs moderately deep, impunctate. Intervals impunctate, slightly and stones. convex. Interval 3 without setiferous puncture behind mid- Remarks. This taxon is morphologically close to K. dle. Aedeagus (Fig. 62). Lateral view: as for genus; moder- embersoni but it is geographically isolated from it, being ately arcuate; apex broadly triangular. Dorsal view: as for found south of the Aupouri Peninsula whereas K. embersoni genus; apical disc present, wide, with convergent sides. is restricted to the tip of the Peninsula. This species is Material examined. 12 specimens, including types named after Barbara I. P. Barratt (AgResearch, Mosgiel) (AMNZ, MONZ, NZAC). for her contribution to the building of important reference collections of New Zealand carabids. Geographic distribution (Map p. 150). North Island: ND–Kohuronaki. North Cape. Pandora. Sandy Bay. Tapotupotu Bay. E Kupeharpalus embersoni new species Ecology. Lowland. Wet forests (broadleaf, podocarp): along Figures 62, 145, 201; Map p. 150 streams. Shaded ground; soil covered with dead leaves. Kupeharpalus embersoni Larochelle & Larivière, new spe- Nocturnal; sheltering during the day in leaf litter (mostly) cies. Holotype: male (NZAC) labelled “NEW ZEALAND ND Te Paki Res, Sandy Bay 16.II. 1995 Larivière, and under logs. Biology. Seasonality: November–Febru- Larochelle (typed) / Streambank; taraire-kauri for; shaded ary, April. Tenerals: February, April. Occasionally infested wet soil, with rich leaf-litter (typed) / HOLOTYPE [male with mites. Dispersal power. Elytra fused along suture. symbol] Kupeharpalus embersoni Larochelle & Larivière, Subapterous. Moderate runner. Collecting techniques. 2004 (red label; typed).” Paratypes: 1 male (MONZ), 1 Raking leaf litter, turning logs, pitfall trapping. female (NZAC) from same locality as holotype, bearing blue paratype labels. Remarks. This species is named after Rowan M. Emberson (Lincoln University, Lincoln) for his contribution to the Description. Body length: 6.0–6.5 mm. Strongly convex. building of important reference collections of New Zea- Piceous brown; antennae, palpi, and legs yellowish. Gen- land carabids. See also Remarks under K. barrattae. erally glabrous and smooth. Microsculpture isodiametric on head, moderately transverse on thorax, very transverse (with microlines) on elytra. Head and pronotum moder- E ately shiny; elytra less shiny, iridescent; without metallic Kupeharpalus johnsi new species lustre. Head. Moderately large, narrower across eyes than Figures 63, 104, 146, 202; Map p. 150 pronotal apex; flat anteriorly, slightly convex posteriorly. Kupeharpalus johnsi Larochelle & Larivière, new species. Labrum strongly transverse; apex slightly emarginate. Eyes Holotype: male (NZAC) labelled “Kara, Whangarei widely separated from buccal fissure ventrally (by about 16.7.27 (hand-written) / coll. E. Fairburn (hand-written) × / HOLOTYPE [male symbol] Kupeharpalus johnsi 1.5 maximum width of antennal scape). Frons with clypeo- Larochelle & Larivière, 2004 (red label; typed).” ocular prolongations incomplete toward eyes. Antennae Paratypes: 9 males (1 MONZ, 4 NZAC, 4 CMNZ) and 1 moderately long, reaching pronotal base; antennal scape female (CMNZ) from the same locality as the holotype, about 2× longer than its maximum width. Paraglossae longer bearing blue paratype labels. 60 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Description. Body length: 5.0–6.0 mm. Strongly convex. congeners (see characters in key to species). It is named Forebody (head and thorax) dark brown; elytra, labrum, after Peter M. Johns (Canterbury Museum, Christchurch) antennae, palpi, and legs rufous. Generally glabrous and for his special help in our entomological studies and for his punctate. Microsculpture absent on forebody, absent or contribution to the building of important reference collec- very transverse (with microlines) on elytra. Very shiny, tions of New Zealand carabids. without metallic lustre. Head. Moderately large, narrower across eyes than pronotal apex; excavated anteriorly, N slightly convex posteriorly. Labrum slightly transverse, Genus Lecanomerus Chaudoir, 1850 Lecanomerus Chaudoir, 1850: 446. Type species: almost square; apex straight or slightly emarginate medi- Lecanomerus insidiosus Chaudoir, 1850, by monotypy. ally. Eyes widely separated from buccal fissure ventrally Thenarotes Bates,1878a: 320. Type species: Thenarotes × (by about 2 maximum width of antennal scape). Frons tasmanicus Bates, 1878a, by monotypy. Synonymised with clypeo-ocular prolongations complete. Antennae by Sloane, 1920: 137. moderately long, reaching elytral base; antennal scape about Odontagonum Darlington, 1956: 8. Type species: 2× longer than its maximum width. Paraglossae as long as Odontagonum nigrum Darlington, 1956, by monotypy. ligula. Penultimate segment of labial palpi with 2 long setae Synonymised by Moore, in Moore et al., 1987: 225. and 1 short seta on anterior margin. Thorax. Pronotum Description. Body length: 3.2–10.0 mm. Forebody (head (Fig. 146) very transverse, widest before middle; sides and thorax) without sparse setiferous micropores dorsally. converging toward base, slightly sinuate; base straight, Head. Mandibles short or moderately long, slightly or moderately narrower than elytral base; apex slightly con- strongly curved forward, acute apically. Labrum strongly cave; lateral depressions widening posteriorly; anterior bead or moderately transverse (atriceps, insignitus, marrisi); incomplete medially; posterior bead complete; anterior apex straight or slightly emarginate medially. Eyes moder- angles moderately developed, obtuse; posterior angles ately large, convex, narrowly separated from buccal fissure strongly developed, subrectangular; basal foveae deep, nar- ventrally (by 0.3–0.5× maximum width of antennal scape), row; punctuation strongly developed (basally). Apex of or reaching buccal fissure (marrisi, sharpi, verticalis). prosternal lobe with 10–20 short setae (without long se- Tempora not inflated. Frons with clypeo-ocular tae). Metepisterna wider than long. Legs. Male mesotarsi prolongations complete or incomplete toward eyes unmodified, neither dilated laterally nor spongily pubes- (insignitus, marrisi). Antennal pubescence starting on cent ventrally. Metatarsomere 5 pubescent (with 5–6 se- antennomere 3 or 2 (atriceps, vestigialis). Mentum with a tae) ventrally. Elytra. Widest behind middle. Shoulders tooth medially, moderately shorter, much shorter (atriceps) strongly developed, rounded, without a tooth. Subapical or about as long as lateral lobes (marrisi). Mentum and sinuations feeble. Sutural apices angulate. Scutellar striole submentum separated by complete transverse suture. absent or present. Interneurs moderately deep, impunctate. Paraglossae longer than or as long as ligula (vestigialis). Intervals impunctate, slightly convex. Interval 3 without Palpi with last segment fusiform or cylindrical (insignitus), setiferous puncture behind middle. Aedeagus (Fig. 63). not truncate apically, sparsely pubescent (with very short Lateral view: as for genus; strongly arcuate; apex narrowly or moderately long setae), or glabrous (insignitus, marrisi); pointed; main shaft much narrower than in other penultimate segment of labial palpi bisetose on anterior Kupeharpalus species. Dorsal view: as for genus; apical margin. Thorax. Pronotum transverse; base straight, disc absent. emarginate (insignitus, marrisi) or convex, moderately nar- Material examined. 36 specimens, including types rower than or as wide as elytral base; lateral beads com- (AMNZ, CMNZ, JNNZ, MONZ, NZAC). plete; anterior bead incomplete medially or complete (insignitus); posterior bead incomplete medially. Scutellum Geographic distribution (Map p. 150). North Island: visible. Apex of prosternal lobe glabrous. Legs. Metafemora AK–Matakana. BP–Kaimai Summit. CL– Coromandel with 2 long setae on posterior margin. Male protarsi di- Range. Maumaupaki Track. Tapu-Coroglen Road. ND– lated laterally and spongily pubescent ventrally. Male Hikurangi. Kara. Parakao. Tangihua Range. mesotarsi dilated laterally and spongily pubescent ventrally Ecology. Lowland, montane. A moist broadleaf forest, along (with spongy pubescence uniformly distributed, contrary a stream, with shaded ground covered with thick leaf litter. to Kupeharpalus) or unmodified (marrisi). Segment 4 of Gregarious. Biology. Seasonality: September–March, July. protarsi and mesotarsi of both sexes without membranous Tenerals: March. Occasionally infested with mites. Dis- laminae. Tarsi pubescent (with numerous setae or only a persal power. Elytra fused along suture. Subapterous. few (verticalis)) or glabrous (insignitus, sharpi) dorsally; Moderate runner. Collecting techniques. Pitfall trapping. metatarsomere 5 pubescent (with 4–8 setae) ventrally; Remarks. This taxon is morphologically isolated from its metatarsomere 1 as long as or shorter than (atriceps, Fauna of New Zealand 53 61 latimanus, marrisi) metatarsomeres 2+3. Elytra. Interneurs 3(1) Pronotum (Fig. 147): reddish brown, contrasting with complete. Rows of setiferous punctures absent on inter- dark head and elytra (Fig. 203); base much narrower vals 3, 5, and 7, and on interneur 2. Umbilicate setiferous than apex. Slender body, somewhat parallel-sided (Fig. series of interval 9 separated into two major groups, with 203). Mentum with medial tooth very small, much posterior group further divided into two subgroups, or, shorter than lateral lobes (Fig. 17). Body length 3.5 posterior group continuous (insignitus). Abdomen. mm or less ...... (p. 62)... atriceps (Macleay) Ventrites 2+3 of male without a setiferous fovea. Ventrites — Pronotum: not reddish brown; base about as wide as 5+6 of both sexes without short setae, with paired ambu- apex. Subovate (somewhat egg-shaped) body. Mentum latory setae only. Aedeagus. Lateral view: slightly to with medial tooth moderately shorter (Fig. 14) than or strongly arcuate. Dorsal view: symmetrical (with ostium about as long as lateral lobes (Fig. 16). Body length 4.0 dorsal, not deflected laterally); dorsal membranous area mm or more (usually over 5 mm) ...... 4 wide, extending almost to basal bulb; apical disc absent or 4(3) Pronotum (Fig. 152) sinuate in front of subrectangular present. Internal sac armed or unarmed. posterior angles. Mentum with medial tooth as long as Geographic distribution. New Guinea, New Caledonia, lateral lobes (Fig. 16). Male mesotarsi unmodified Australia (including Tasmania), New Zealand. (neither dilated laterally (Fig. 208) nor spongily References. Hudson, 1934: 176 (list); Noonan, 1976: 7 pubescent ventrally) [South Island: Banks Peninsula] (taxonomy); Larochelle & Larivière, 2001: 118–121 (cata- ...... (p. 66)... marrisi new species logue). — Pronotum not sinuate in front of rounded posterior Remarks. Sloane (1920: 137) incorrectly synonymised angles. Mentum with medial tooth moderately shorter Lecanomerus with Nemaglossa Solier, 1849, which is a than lateral lobes (Fig. 14). Male mesotarsi dilated valid genus restricted to Chile (Noonan, 1976: 7). laterally (Fig. 205) and spongily pubescent ventrally Lecanomerus is a highly variable genus which currently (Fig. 12) ...... 5 includes species with the following combination of charac- 5(4) Elytra fused along suture; membranous wings vestigial ters: eyes reaching buccal fissure or narrowly separated (reduced to wing buds) ...... 6 from it; penultimate segment of labial palpi bisetose; apex — Elytra not fused along suture; membranous wings fully of prosternal lobe glabrous; clypeo-ocular prolongations developed ...... 7 complete (incomplete toward eyes in insignitus, marrisi); umbilicate setiferous series of interval 9 with posterior 6(5) Pronotum (Fig. 148): base emarginate; basal foveae group divided into 2 subgroups (continuous in insignitus). shallow, weakly and finely punctate. Frons with Australian Lecanomerus species are diverse and in great clypeo-ocular prolongations incomplete toward eyes need of revision supported by a cladistic analysis; only (Fig. 105). [North Island: ND] ...... then will we be able to define the taxonomic limits of this ...... (p. 63)... insignitus Broun genus and the generic placement of currently described — Pronotum (Fig. 149): base rather straight; basal foveae species in New Zealand and the rest of Australasia. deep, strongly and coarsely punctate. Frons with clypeo-ocular prolongations complete (Fig. 1). [South Island]...... (p. 63)... obesulus Bates Key to species of Lecanomerus 7(5) Elytra very iridescent, with subapical sinuations 1 Elytra with sides broadly yellowish (Fig. 206–207) . 2 strongly developed (Fig. 30). Pronotum (Fig. 153) — Elytra with sides not broadly yellowish (Fig. 203–205, widest before middle. Appendages entirely pale reddish. 208–210) ...... 3 Body slightly convex; length 4.7 mm or more ...... (p. 67)... verticalis (Erichson) 2(1) Pronotum (Fig. 151): sides slightly sinuate before strongly developed, acute posterior angles with — Elytra slightly iridescent, with subapical sinuations prominent tooth. Pale outline of elytra somewhat weakly developed (Fig. 29). Pronotum (Fig. 154) jagged (Fig. 207). [TH, North Island]...... widest about middle. Appendages partly pale reddish ...... (p. 65)... sharpi (Csiki) (parts of antennae, tibiae, and tarsi dark brown). Body strongly convex; length 4.3 mm or less ...... — Pronotum (Fig. 150): sides not sinuate before moderately ...... (p. 67)... vestigialis (Erichson) developed, obtusely rounded posterior angles, without tooth. Pale outline of elytra more regular, not jagged (Fig. 206). [CH, South Island] ...... (p. 64)... latimanus Bates 62 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Lecanomerus atriceps (Macleay, 1871) A ture behind middle. Umbilicate setiferous series of interval Figures 64, 147, 203; Map p. 150 9 with posterior group divided into 2 subgroups. Aedeagus Trechus atriceps Macleay, 1871: 113. Type locality: Gayndah, (Fig. 64). Lateral view: as for genus; strongly arcuate; apex Queensland, Australia. broadly triangular. Dorsal view: as for genus; apical disc Thenarotes atriceps: Blackburn, 1892: 97. absent. Internal sac unarmed. Stouter and smaller than in Nemaglossa atriceps: Sloane, 1920: 137. other Lecanomerus species. Nemaglossa (Thenarotes) atriceps: Pilgrim, 1963: 844. Lecanomerus atriceps: Moore et al., 1987: 225. Material examined. 89 non-type specimens (AMNZ, Acupalpus javanus Jedli…ka, 1964: 311. Type locality: BMNH, CMNZ, ITNZ, JNNZ, LUNZ, MONZ, UCNZ). Puntjak-Pass, W. Java (probably mislabelled (Kataev, Geographic distribution (Map p. 150). North Island: 2002b). Synonymised by Kataev, 2002b: 722. AK, BP, CL, ND, WI, WN. South Island: MC, NN, SD. Description. Body length: 3.2–3.5 mm. Slender, more or Adventive. Extralimital range: Australia (mainland). First less parallel-sided (more so than other Lecanomerus spe- New Zealand records: Swanson, AK, 1916 (NZAC); near cies). Slightly convex. Head black; pronotum reddish brown; Auckland, AK, 1916 (Pilgrim, 1963: 845); Port Waikato, elytra dark brown; elytral margins and sutures red; WO, 1958 (May, 1963: 192). Well established. antennomeres 1+2, palpi, and legs yellowish; antennomeres Ecology. Lowland. Borders of eutrophic marshes and 3–11 blackish (contrary to other Lecanomerus species). ponds, and slowly running rivers. Wet pastures, paddocks, Generally glabrous and smooth. Microsculpture vestigial and gardens. Open ground; soil inundated, covered with on forebody (isodiametric on head, moderately transverse dense tall vegetation (e.g., Cyperaceae, Typha) or alligator on pronotum), and very transverse (with microlines) on weed mats, or, when non-inundated, soft, wet, muddy or elytra. Shiny, without metallic lustre; elytra very irides- clayey, covered with similar vegetation, leaf litter or nikau cent. Head. Moderately large, narrower across eyes than sheaths. Occasional in caves. Nocturnal; sheltering during pronotal apex; flat anteriorly, slightly convex posteriorly. the day in floating plant mats, at the base of plants, in leaf Mandibles short, strongly curved forward. Labrum mod- litter, under nikau sheaths, stones, or clay cakes. Gregari- erately transverse; apex slightly emarginate medially. Eyes ous. Biology. Seasonality: throughout the year. Tenerals: narrowly separated from buccal fissure ventrally (by about December (rarely), April, June–July. Predacious (Moore × 0.3 maximum width of antennal scape). Frons with clypeo- et al., 1987: 225). Occasionally infested with mites. Dis- ocular prolongations complete. Antennae moderately long, persal power. Elytra free along suture. Macropterous. × reaching pronotal base; antennal scape about 2 longer Frequent flier. Occasional in seashore drift material, which than its maximum width; pubescence starting from apical indicates flight ability. Moderate runner. Occasional climber 1/2 of antennomere 2. Mentum with a very short tooth (on plants). Good burrower. Clearly effective as a colonist. × medially, much shorter than lateral lobes (only about 0.3 Favoured by human activities. Collecting techniques. as long). Paraglossae longer than ligula. Palpi with last Treading plant stems or carpets into the water, turning segment fusiform, with sparse, moderately long pubes- plant debris, sweeping vegetation, light trapping, pitfall cence. Thorax. Pronotum (Fig. 147) moderately trans- trapping. verse, widest before middle; sides converging toward base, References. May, 1963: 192 (distribution, ecology); Pil- not sinuate; base slightly convex, moderately narrower than grim, 1963: 844–845 (distribution); Johns, 1986: 31 (dis- elytral base, much narrower than pronotal apex (contrary tribution, ecology); Moore et al., 1987: 225 (distribution, to other Lecanomerus species); apex straight; lateral de- ecology, biology, dispersal power); Townsend, 1994: 9, pressions absent; anterior angles slightly developed, ob- 11–12 (distribution, ecology); Larochelle & Larivière, 2001: tuse; posterior angles moderately developed, obtusely 119 (taxonomy, distribution, ecology, biology, dispersal rounded; basal foveae shallow, wide; punctuation strongly power). developed and coarse (in basal foveae). Metepisterna longer than wide. Legs. Male mesotarsi dilated laterally and Remarks. The reddish brown pronotum contrasting with spongily pubescent ventrally. Tarsi pubescent (with nu- the dark head and elytra, and the body shape make this merous setae) dorsally; metatarsomere 5 pubescent (with species superficially similar to Euthenarus bicolor. Both 6 setae) ventrally; metatarsomere 1 shorter than species have often been confused in the past, but they are metatarsomeres 2+3. Elytra. Widest about middle. Shoul- easily distinguished based on male tarsal characters (tarsi ders strongly developed, rounded, without a tooth. Sub- spongily pubescent ventrally in L. atriceps, biseriately apical sinuations feeble. Sutural apices angulate. Scutellar pubescent in E. bicolor) and ventral pubescence (ventrites striole absent. Interneurs complete, shallow, impunctate. 5+6 without setae and setiferous fovea of ventrites 2+3 Intervals impunctate, flat. Interval 3 with a setiferous punc- absent in L. atriceps). Fauna of New Zealand 53 63

Lecanomerus insignitus Broun, 1880 E metatarsomeres 2+3. Metatarsal claws of male 2× larger Figures 65, 105, 148, 204; Map p. 150 than pro- and mesotarsal claws (about as wide in other Lecanomerus insignitus Broun, 1880: 47. Holotype: male Lecanomerus species). Elytra. Widest about middle. Shoul- (BMNH) labelled “Type (circular red-bordered label; ders strongly developed, rounded, without a tooth. Sub- typed) / 96. (hand-written) / Parua. (hand-written) / New apical sinuations moderate. Sutural apices angulate. Scutellar Zeal. Broun Coll. Brit. Mus. 1922-482 (typed) / striole absent. Interneurs moderately deep, impunctate. Lecanomerus insignitus (hand-written).” Perfect condi- Intervals impunctate, slightly convex. Interval 3 with a tion. Lecanomerus fallax Broun, 1880: 48. Lectotype (here desig- setiferous puncture behind middle. Umbilicate setiferous nated): male (BMNH) labelled “[male symbol] (hand- series of interval 9 with posterior group continuous. written) / Type (circular red-bordered label; typed) / 99. Aedeagus (Fig. 65). Lateral view: as for genus; slightly (hand-written) / Parua. (hand-written) / Lecanomerus arcuate; apex truncate. Dorsal view: as for genus; apical fallax (hand-written) / LECTOTYPE Lecanomerus fallax disc present, transverse. Internal sac armed. Broun, 1880 designated by Larochelle & Larivière 2004 Material examined. 35 specimens, including types (red label; typed).” Perfect condition. Paralectotypes: 1 male and 1 female (BMNH) bearing blue paralectotype (AMNZ, BMNH, CMNZ, FMNH, JNNZ, LUNZ, labels. New synonym. NZAC). Nemaglossa fallax: Sloane, 1920: 137. Geographic distribution (Map p. 150). North Island: ND. Nemaglossa insignita: Sloane, 1920: 137. Lecanomerus fallax: Hudson, 1934: 176. Ecology. Lowland. Wet forests (broadleaf, podocarp): along Lecanomerus insignitus: Hudson, 1934: 176. streams. Shaded ground; soil covered with dead leaves. Nocturnal; sheltering during the day in leaf litter and under Description. Body length: 5.0–10.0 mm. Strongly con- logs. Gregarious. Biology. Seasonality: October–April, vex. Brownish black; pronotal margins, elytral margins, July–August. Dispersal power. Elytra fused along suture. elytral apex, and appendages brownish red. Generally gla- Subapterous. Moderate runner. Collecting techniques. brous and smooth. Microsculpture isodiametric on head, Raking leaf litter, turning logs, pitfall trapping. moderately transverse on pronotum, and very transverse (with microlines) on elytra. Moderately shiny, without Reference. Larochelle & Larivière, 2001: 119 (including metallic lustre; elytra very iridescent. Head. Rather small, fallax; taxonomy, distribution, ecology, biology, dispersal much narrower across eyes than pronotal apex; flat power). anteriorly, slightly convex posteriorly. Mandibles moder- Remarks. Broun described Lecanomerus fallax from 4 ately elongate, slightly curved forward. Labrum moder- specimens, only 3 of which could be located in the Natural ately transverse, almost quadrate; apex straight or slightly History Museum, London (BMNH). One of these speci- emarginate medially. Eyes narrowly separated from buccal mens is designated as lectotype to preserve stability of fissure ventrally (by about 0.7× maximum width of antennal nomenclature in the future. This species may appear su- scape). Frons with clypeo-ocular prolongations incom- perficially similar to Kupeharpalus barrattae and K. plete toward eyes. Antennae moderately long, reaching embersoni, but the glabrous apex of the prosternal lobe, pronotal base; antennal scape about 2× longer than its maxi- the bisetose penultimate segments of the labial palpi, and mum width; pubescence starting from basal 1/3 of the eyes narrowly separated from the buccal fissure antennomere 3. Mentum with a tooth medially, moder- ventrally, clearly place it in Lecanomerus. The body size ately shorter than lateral lobes. Paraglossae longer than and external morphology of L. insignitus are highly vari- ligula. Palpi with last segment cylindrical, glabrous. Tho- able but the configuration of the male genitalia is constant rax. Pronotum (Fig. 148) very transverse, widest about throughout the geographical range of this species. middle; sides converging toward base, not sinuate; base emarginate, as wide as elytral base; anterior bead complete (contrary to other Lecanomerus species); apex slightly Lecanomerus obesulus Bates, 1878 E concave; lateral depressions widening posteriorly; anterior Figures 66, 149, 205; Map p. 151 angles moderately developed, rounded; posterior angles Lecanomerus obesulus Bates, 1878c: 23. Lectotype (here strongly developed, broadly rounded; basal foveae shal- designated): male (MNHN) labelled “ W. Coast S.I., N.Z. low, ill-defined; punctuation feebly developed basally. (hand-written) / Lecanomerus obesulus Bates (hand-writ- ten) / LECTOTYPE Lecanomerus obesulus Bates, 1878 Metepisterna as wide as long. Legs. Male protarsi 3x wider designated by Larochelle & Larivière 2004 (red label; than mesotarsi (contrary to other Lecanomerus species); typed).” Perfect condition. Paralectotypes: 1 male and 1 male mesotarsi dilated laterally and spongily pubescent female (MNHN) bearing blue paralectotype labels. ventrally. Tarsi glabrous dorsally; metatarsomere 5 pubes- Nemaglossa obesula: Sloane, 1920: 137. cent (with 4 setae) ventrally; metatarsomere 1 as long as Lecanomerus obesulus: Hudson, 1934: 176. 64 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Description. Body length: 5.0–5.3 mm. Moderately con- Moderate runner. Collecting techniques. Turning moss vex. Black; margins of pronotum and elytra narrowly red- carpets, stones, and logs. dish; antennomeres 1+2 and basal 1/2 of tibiae reddish; References. Townsend, 1997: 17 (distribution); Larochelle femora blackish. Generally glabrous and smooth. & Larivière, 2001: 119–120 (taxonomy, distribution, ecol- Microsculpture vestigial on forebody (head and thorax), ogy, biology, dispersal power). isodiametric on head, moderately transverse on pronotum, Remarks. Bates’ original description was based on “nu- very transverse (with microlines) on elytra. Shiny, with merous examples” from “West Coast, S. [=South] Island.” slight brassy metallic lustre on elytra; moderately irides- Three syntypes were obtained from the Bates collection in cent. Head. Moderately large, narrower across eyes than Paris (MNHN), one of which (a male specimen) bears a pronotal apex; flat anteriorly, slightly convex posteriorly. determination label written by Bates; this specimen is here Mandibles short, strongly curved forward. Labrum selected as lectotype to preserve stability of nomenclature strongly transverse; apex slightly emarginate medially. Eyes in the future. See under L. latimanus. narrowly separated from buccal fissure ventrally (by about 0.5× maximum width of antennal scape). Frons with clypeo- ocular prolongations complete. Antennae moderately long, E × Lecanomerus latimanus Bates, 1874 reaching pronotal base; antennal scape about 2 longer Figures 67, 150, 206; Map p. 150 than its maximum width; pubescence starting from basal Lecanomerus latimanus Bates, 1874: 271 (redescribed in third of antennomere 3. Mentum with a tooth medially, 1875: 309). Holotype: male (MNHN) labelled “New moderately shorter than lateral lobes. Paraglossae longer Zealand. (hand-written) / Lecanomerus latimanus Bates than ligula. Palpi with last segment fusiform, with sparse, (hand-written) / HOLOTYPE [male symbol] very short pubescence. Thorax. Pronotum (Fig. 149) very Lecanomerus latimanus Bates, 1874 (typed; red label).” transverse, widest before middle; sides converging toward Good condition. base, not sinuate; base rather straight, moderately narrower Lecanomerus fuliginosus Broun, 1880: 48. Holotype: male (BMNH) labelled “Type (circular red-bordered label; than elytral base; apex concave; lateral depressions absent; typed) / 98. (typed) / [male symbol] (handwritten) / Taieri anterior angles moderately developed, obtusely rounded; (typed) / New Zeal. Broun Coll. Brit. Mus. 1922-482. posterior angles moderately developed, broadly rounded; (typed) / Lecanomerus fuliginosus (hand-written).” Per- basal foveae deep, wide; punctuation strongly developed fect condition. New synonym. and coarse basally. Metepisterna longer than wide. Legs. Lecanomerus pallipes Broun, 1894: 379. Holotype: male Male mesotarsi dilated laterally and spongily pubescent (BMNH) labelled “Type (circular red-bordered label; ventrally. Tarsi pubescent (with numerous setae) dorsally; typed) / 2623. (handwritten) / [male symbol] (hand-writ- metatarsomere 5 pubescent (with 6 setae) ventrally; ten) / Canterbury (typed) / New Zealand. Broun Coll. metatarsomere 1 as long as metatarsomeres 2+3. Elytra. Brit. Mus. 1922-482. (typed) / Lecanomerus pallipes. (hand-written).” Perfect condition. New synonym. Widest about middle. Shoulders strongly developed, Lecanomerus incertus Broun, 1914b: 151. Holotype: male rounded, without a tooth. Subapical sinuations feeble. (BMNH) labelled “Type (circular red-bordered label; Sutural apices rounded. Scutellar striole absent. Interneurs typed) / 3512 [male symbol] (hand-written) / New Zeal. shallow, impunctate. Intervals impunctate, flat. Interval 3 Broun Coll. Brit. Mus. 1922-482 (typed) / Mount Hutt without setiferous puncture. Umbilicate setiferous series Decr. 1911. (hand-written) / Lecanomerus incertus. [male of interval 9 with posterior group divided into two sub- symbol]. (hand-written).” Perfect condition. New syno- groups.Aedeagus (Fig. 66). Lateral view: as for genus; nym. strongly arcuate; apex acute. Dorsal view: as for genus; Nemaglossa latimana: Sloane, 1920: 137. Nemaglossa fuliginosa: Sloane, 1920: 137. apical disc absent. Internal sac armed. Nemaglossa pallipes: Sloane, 1920: 137 Material examined. 83 specimens, including types Nemaglossa incerta: Sloane, 1920: 137. (AMNZ, BBNZ, CMNZ, ITNZ, LUNZ, MNHN, Lecanomerus incertus: Hudson, 1934: 176. MONZ, NZAC, PHNZ, UCNZ). Lecanomerus latimanus: Hudson, 1934: 176. Lecanomerus fuliginosus: Hudson, 1934: 176. Geographic distribution (Map p. 151). South Island: Lecanomerus pallipes: Hudson, 1934: 176. BR, CO, FD, MB, MC, NC, NN, OL, SD, SL, WD. Description. Body length: 4.7–5.9 mm. Slightly convex. Ecology. Lowland, montane. Forests (beech, podocarp, Head and pronotum brownish testaceous; elytra blackish broadleaf) and scrublands (bog pine). Shaded ground. Noc- with sides broadly yellowish; appendages yellowish. Gen- turnal; sheltering during the day in moss carpets, under erally glabrous and smooth. Microsculpture strongly de- stones and logs. Gregarious. Biology. Seasonality: Sep- veloped, granulate on head, isodiametric on pronotum, and tember–February, April–June. Tenerals: February–March. moderately transverse on elytra. Dull; elytra iridescent. Dispersal power. Elytra fused along suture. Subapterous. Head. Moderately large, narrower across eyes than pronotal Fauna of New Zealand 53 65 apex; flat anteriorly, slightly convex posteriorly. Mandi- fuliginosus; distribution); Emberson, 1998: 30 (as bles short, strongly curved forward. Labrum strongly trans- fuliginosus; distribution, ecology, biology); Larochelle & verse; apex slightly emarginate medially. Eyes narrowly Larivière, 2001: 119–120 (including fuliginosus, incertus, separated from buccal fissure ventrally (by about 0.5× pallipes; taxonomy, distribution, ecology, biology, disper- maximum width of antennal scape). Frons with clypeo- sal power. ocular prolongations complete. Antennae moderately long, Remarks. Bates’ original description of Lecanomerus × reaching pronotal base; antennal scape about 2 longer latimanus was based on “one example from New Zea- than its maximum width; pubescence starting from basal land.” Five specimens were obtained from the Bates col- 1/3 of antennomere 3. Mentum with a tooth medially, mod- lection in Paris (MNHN), only one of which, a male, erately shorter than lateral lobes. Paraglossae longer than matches data in the original description. The authors think ligula. Palpi with last segment fusiform, with sparse, very that this is the specimen used by him for the description, short pubescence. Thorax. Pronotum (Fig. 150) very trans- although not labelled as such by Bates; a red holotype label verse, widest before middle; sides converging toward base, has thus been added to the MNHN specimen. In some not sinuate; base straight, moderately narrower than elytral populations the elytral sides may be more narrowly yel- base; apex concave; lateral depressions widening lowish in basal half, but the configuration of the male geni- posteriorly; anterior angles strongly developed, obtusely talia is stable. This species resembles L. sharpi in its gen- rounded; posterior angles moderately developed, obtusely eral shape and broadly yellowish elytral sides. Lecanomerus rounded; basal foveae shallow, narrow, linear; punctuation latimanus and L. obesulus are the most commonly encoun- feebly developed basally. Metepisterna longer than wide. tered Lecanomerus species on the South Island. Legs. Male mesotarsi dilated laterally and spongily pu- bescent ventrally. Tarsi pubescent (with numerous setae) dorsally; metatarsomere 5 pubescent (with 6 setae) Lecanomerus sharpi (Csiki, 1932) E ventrally; metatarsomere 1 shorter than metatarsomeres Figures 68, 151, 207; Map p. 151 2+3. Elytra. Widest about middle. Shoulders strongly de- Lecanomerus marginatus Sharp, 1883: 25. Holotype: male veloped, rounded, without a tooth. Subapical sinuations (BMNH) labelled “Lecanomerus marginatus type D.S. d feeble. Sutural apices angulate. Scutellar striole absent or Auckland. N. Zeal . (hand-written) / Type H.T. (circular red-bordered label; typed) / Sharp Coll. 1905-313 present. Interneurs shallow, impunctate. Intervals (typed).” Perfect condition. Primary homonym of impunctate, flat. Interval 3 without setiferous puncture Lecanomerus marginatus Reed, 1874. behind middle. Umbilicate setiferous series of interval 9 Nemaglossa sharpi Csiki, 1932: 1059 (replacement name with posterior group divided into two subgroups. Aedeagus for Lecanomerus marginatus Sharp, 1883). (Fig. 67). Lateral view: as for genus; strongly arcuate; apex Nemaglossa marginata: Sloane, 1920: 137. acute. Dorsal view: as for genus; apical disc absent. Inter- Lecanomerus sharpi: Noonan, 1976: 7. nal sac armed. Description. Body length: 5.0–6.0 mm. Slightly convex. Material examined. 106 specimens, including types Piceous brown; sides of pronotum and elytra broadly yel- (BMNH, CMNZ, ITNZ, JNNZ, LUNZ, MNHN, NZAC, lowish; antennomere 1, femora, basal 1/2 of tibiae, and OMNZ, PHNZ, UCNZ). tarsi yellowish. Generally glabrous and smooth. Microsculpture isodiametric on head, transverse on Geographic distribution (Map p. 150). South Island: pronotum and elytra. Shiny, without metallic lustre. Head. CO, DN, KA, MC, SC, SL. Offshore Islands: CH. Moderately large, narrower across eyes than pronotal apex; Ecology. Lowland, upland, subalpine, alpine. Tussock flat anteriorly, slightly convex posteriorly. Mandibles short, grasslands (mostly), herbfields, farmlands, gardens, river strongly curved forward. Labrum strongly transverse; apex banks, screes, forests (beech, broadleaf, podocarp). Open slightly emarginate medially. Eyes reaching buccal fissure and shaded ground; soil dry or moist, covered with grass or ventrally. Frons with clypeo-ocular prolongations com- dead leaves. Nocturnal; sheltering during the day at the plete. Antennae moderately long, reaching pronotal base; base of tussock-plants, in moss, mat plants, plant debris, antennal scape about 2× longer than its maximum width; under stones, under and in logs, and in leaf litter. Biology. pubescence starting from basal 1/2 of antennomere 3. Men- Seasonality: September–June. Tenerals: May. Dispersal tum with a tooth medially, moderately shorter than lateral power. Elytra fused along suture. Subapterous. Moderate lobes. Paraglossae longer than ligula. Palpi with last seg- runner. Collecting techniques. Inspecting the base of ment fusiform, with sparse, very short pubescence. Tho- tussock-plants, turning stones, raking leaf litter, pitfall trap- rax. Pronotum (Fig. 151) very transverse, widest before ping. middle; sides converging toward base, slightly sinuate; base References. Barratt & Patrick, 1987: 82 (as fuliginosus; rather straight, moderately narrower than elytral base; apex distribution, ecology, biology); Townsend, 1997: 17 (as concave; lateral depressions widening posteriorly; anterior 66 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) angles moderately developed, rounded; posterior angles Dark brown; labrum and clypeus reddish; appendages, strongly developed, acute, with a prominent tooth (con- pronotal margins, sutures, epipleura, and subapical mar- trary to other Lecanomerus species); basal foveae shallow, gins of elytra brownish red. Generally glabrous and punc- wide; punctuation feebly developed basally. Metepisterna tate. Microsculpture absent on head, very transverse (with longer than wide. Legs. Male mesotarsi dilated laterally microlines) on pronotum and elytra. Shiny, without metal- and spongily pubescent ventrally. Tarsi glabrous dorsally; lic lustre; pronotum and elytra iridescent. Head. Moder- metatarsomere 5 pubescent (with 4 setae) ventrally; ately large, narrower across eyes than pronotal apex; exca- metatarsomere 1 as long as metatarsomeres 2+3. Elytra. vated anteriorly, slightly convex posteriorly. Mandibles Widest about middle. Shoulders strongly developed, moderately long, slightly curved forward. Labrum strongly rounded, without a tooth. Subapical sinuations feeble. transverse; apex slightly emarginate medially. Eyes reach- Sutural apices angulate. Scutellar striole absent or present. ing buccal fissure ventrally. Frons with clypeo-ocular Interneurs shallow, impunctate. Intervals impunctate, flat. prolongations incomplete toward eyes. Antennae moder- Interval 3 without setiferous puncture. Umbilicate ately long, reaching about elytral base; antennal scape about setiferous series of interval 9 with posterior group divided 2× longer than its maximum width; pubescence starting on into 2 subgroups. Aedeagus (Fig. 68). Lateral view: as for basal 1/3 of antennomere 3. Mentum with a tooth medi- genus; slightly arcuate; apex rounded, inflated. Dorsal view: ally, as long as lateral lobes. Paraglossae longer than ligula. as for genus; apical disc present, subtriangular. Internal sac Palpi with last segment fusiform, glabrous. Thorax. armed. Pronotum (Fig. 152) transverse, widest before middle; sides Material examined. 327 specimens, including type converging toward base, slightly sinuate; base slightly (AMNZ, CMNZ, FMNH, ITNZ, JNNZ, LUNZ, MONZ, emarginate, moderately narrower than elytral base; apex NZAC, OMNZ, UCNZ). concave; lateral depressions widening posteriorly; anterior angles strongly developed, obtusely rounded; posterior Geographic distribution (Map p. 151). North Island: angles strongly developed, subrectangular; basal foveae AK, BP, CL, GB, HB, ND, RI, TK, TO, WA, WI, WN, shallow, wide; punctuation strongly developed, fine. WO. Offshore Islands: TH. Metepisterna longer than wide. Legs. Male mesotarsi un- Ecology. Lowland. Wet forests (broadleaf, podocarp) and modified, neither dilated laterally nor spongily pubescent swamp forests: along streams. Shaded ground; soil covered ventrally. Protarsomeres 1–4 pubescent; meso- and with thick leaf litter. Nocturnal; sheltering during the day metatarsomeres 1–4 glabrous; pro-, meso-, and in leaf litter and under stones. Gregarious. Biology. metatarsomeres 5 with 5–6 setae dorsally. Metatarsomere Seasonality: throughout the year. Occasionally infested 5 pubescent (with 5–6 setae) ventrally; metatarsomere 1 with mites. Defense mechanism: feigns death when dis- short, only about as long as metatarsomere 2. Elytra. Wid- turbed. Dispersal power. Elytra fused (brachypterous) or est about middle. Shoulders strongly developed, rounded, free (macropterous) along suture. Moderate runner. Col- without a tooth. Sides rounded. Subapical sinuations fee- lecting techniques. Pitfall trapping, raking leaf litter, bait- ble. Sutural apices rounded. Scutellar striole present. ing pitfall traps, using yellow pan traps. Interneurs shallow, impunctate. Intervals impunctate, flat. References. Johns, 1986: 31 (distribution); Kuschel, 1990: Interval 3 without setiferous puncture behind middle. 24, 40 (distribution, ecology, biology, dispersal power); Umbilicate setiferous series of interval 9 with posterior Larochelle & Larivière, 2001: 120 (taxonomy, distribution, group divided into 2 subgroups. Aedeagus (Fig. 69). Lat- ecology, biology. dispersal power). eral view: as for genus; slightly arcuate; apex obtuse. Dor- Remarks. See under L. latimanus. sal view: as for genus; apical disc absent. Internal sac un- armed. Material examined. 5 specimens, including types (ITNZ, Lecanomerus marrisi new species E NZAC, UCNZ). Figures 69, 106, 152, 208; Map p. 151 Geographic distribution (Map p. 151). South Island: Lecanomerus marrisi Larochelle & Larivière, new species. MC–Banks Peninsula (Head of Kaituna Valley; Peraki– Holotype: male (NZAC) labelled “Peraki-Mt Bossu Rd Mount Bossu Road; Peraki Road Bush). Banks Peninsula 12.12.63 1800' W.P. Thomas (hand- written) / HOLOTYPE [male symbol] Lecanomerus Ecology. Lowland. Habitat unknown. Biology. Seasonality: marrisi Larochelle & Larivière, 2004 (red label; typed).” December, February–March. Tenerals: March. Dispersal Paratypes: 1 male and 2 females (UCNZ) from Peraki Rd power. Elytra fused along suture. Subapterous. Moderate Bush, Banks Peninsula, bearing blue paratype labels. runner (after leg morphology). Description. Body length: 7.0–7.1 mm. Slightly convex. Remarks. The long mentum tooth, anteriorly excavated Fauna of New Zealand 53 67 head, unmodified male mesotarsi, and pronotum shape set mesotarsi dilated laterally and spongily pubescent ventrally. this species apart from its congeners. It is also the only Tarsi pubescent (with a few setae) dorsally; metatarsomere species of Lecanomerus which is endemic to the Banks 5 pubescent (with 6 setae) ventrally; metatarsomere 1 as Peninsula. Lecanomerus marrisi is named after our col- long as metatarsomeres 2+3. Elytra. Widest about middle; league John M. W. Marris (Lincoln University, Lincoln) slightly more elongate than in vestigialis. Shoulders strongly for his special help and encouragement in our entomologi- developed, angulate, with a tooth. Subapical sinuations cal studies. strongly developed. Sutural apices angulate. Scutellar striole absent. Interneurs shallow, impunctate. Intervals

A impunctate, flat. Interval 3 with a setiferous puncture be- Lecanomerus verticalis (Erichson, 1842) hind middle. Umbilicate setiferous series of interval 9 with Figures 70, 153, 209; Map p. 151 posterior group divided into 2 subgroups. Aedeagus (Fig. Harpalus verticalis Erichson, 1842: 126. Type locality: Tas- 70). Lateral view: as for genus; slightly arcuate; basal 1/2 mania, Australia. Lecanomerus insidiosus Chaudoir, 1850: 447. Type locality: much wider than in vestigialis; apex broadly acute. Dorsal Swan River and Melbourne, Australia. Synonymised by view: as for genus; apical disc absent. Internal sac armed. Moore, in Moore et al., 1987: 227. Material examined. 54 non-type specimens (AMNZ, Lecanomerus flavocinctus Blackburn, 1888b: 188. Type lo- ITNZ, JNNZ, MONZ, NZAC). cality: Port Lincoln, South Australia. Synonymised by Moore, in Moore et al., 1987: 227. Geographic distribution (Map p. 151). North Island: Lecanomerus occidentalis Sloane, 1898: 464. Type locality: AK, BP, CL, GB, ND, WI, WN, WO. South Island: NN. Swan River, Darling Range, Pinjarrah, Donnybrook, Mt Adventive. Extralimital range: Australia (including Tasma- Barker, and Albany, Western Australia. Synonymised with nia). First New Zealand records: Okauia, WO, 1922 L. flavocinctus by Sloane, 1911: 835. (NZAC); New Zealand (Moore et al., 1987: 227). Well Lecanomeus verticalis: Sloane, 1911: 835. established. Nemaglossa verticalis: Sloane, 1920: 137. Nemaglossa insidiosa: Sloane, 1920: 137. Ecology. Lowland, upland. Sand dunes, vacant lots, and Nemaglossa flavocincta: Sloane, 1920: 137. gardens. Open ground; dry, sandy soil sparsely vegetated Lecanomerus verticalis: Noonan, 1976: 7. by grass and weeds. Nocturnal; sheltering during the day in Description. Body length: 4.7–6.5 mm. Slightly convex. burrows dug at the base of plants, under pieces of wood, Piceous black; sides of pronotum and elytra narrowly pale and in garden rubbish. Gregarious. Biology. Seasonality: reddish; appendages pale reddish. Generally glabrous and September–June. Tenerals: October–November, January– smooth. Microsculpture strong, isodiametric on head, February. Predacious (Moore et al., 1987: 227). Occasion- moderately transverse on pronotum, and very transverse ally infested with mites and fungi (Laboulbeniales). Dis- (with microlines) on elytra. Dull, without metallic lustre; persal power. Elytra free along suture. Macropterous. elytra very iridescent. Head. Moderately large, narrower Occasional flier. Moderate runner. Good burrower. Clearly across eyes than pronotal apex; flat anteriorly, slightly effective as a colonist. Favoured by human activities. Col- convex posteriorly. Mandibles short, strongly curved for- lecting techniques. Pitfall trapping, digging at the base of ward. Labrum strongly transverse; apex slightly emarginate plants, turning pieces of wood. medially. Eyes reaching buccal fissure ventrally. Frons with References. Moore et al., 1987: 227 (distribution, ecol- clypeo-ocular prolongations complete. Antennae moder- ogy, biology, dispersal power); Townsend, 1994: 9, 11 and ately long, reaching pronotal base; antennal scape about 2× 1997: 17 (distribution, ecology); Larochelle & Larivière, longer than its maximum width; pubescence starting from 2001: 120–121 (taxonomy, distribution, ecology, biology, apex of antennomere 2. Mentum with a tooth medially, dispersal power). moderately shorter than lateral lobes. Palpi with last seg- Remarks. This species is morphologically close to L. ment fusiform, with sparse, very short pubescence. Tho- vestigialis (see characters in key to species). rax. Pronotum (Fig. 153) very transverse, widest before middle (about middle in vestigialis); sides converging to- ward base, not sinuate; base slightly convex, moderately Lecanomerus vestigialis (Erichson, 1842) A narrower than elytral base; apex concave; lateral depres- Figures 71, 107, 154, 210; Map p. 151 sions widening posteriorly; anterior angles moderately de- Harpalus vestigialis Erichson, 1842: 127. Type locality: veloped, rounded; posterior angles moderately developed, Tasmania, Australia. broadly rounded; basal foveae deep, very wide (touching Acupalpus mastersii Macleay, 1871: 104. Type locality: lateral beads); punctuation developed basally, moderately Gayndah, Queensland, Australia. Synonymised by Moore, coarse. Metepisterna longer than wide. Legs. Male in Moore et al., 1987: 227. 68 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Lecanomerus stenopus Broun, 1886: 880. Holotype: female than in verticalis. Shoulders strongly developed, angulate, (BMNH) labelled “Type (circular red-bordered label; with a tooth. Subapical sinuations feebly developed. Sutural typed) / 1565. (hand-written) / New Zealand Broun Coll. apices angulate. Scutellar striole absent or present. r Brit. Mus. 1922-482 (typed) / Howick. Nov . 1885. (hand- Interneurs shallow, impunctate. Intervals impunctate, flat. written) / Lecanomerus stenopus. [female symbol]. (hand- written).” Perfect condition. Synonymised by Moore, in Interval 3 without setiferous puncture behind middle. Moore et al., 1987: 227. Umbilicate setiferous series of interval 9 with posterior Lecanomerus nitidus Blackburn, 1891: 779. Type locality: group divided into two subgroups. Aedeagus (Fig. 71). Victoria, Australia. Synonymised with Nemaglossa Lateral view: as for genus; slightly arcuate; basal 1/2 much mastersi by Sloane, 1920: 137. narrower than in verticalis; apex broadly acute. Dorsal Lecanomerus mastersii: Sloane, 1911: 836. view: as for genus; apical disc absent. Internal sac armed. Lecanomerus labralis Broun, 1914b: 151. Holotype: male (BMNH) labelled “Type (circular red-bordered label; Material examined. 294 specimens, including Broun’s typed) / 3511. [male symbol]. (hand-written) / Epsom. types (AMNZ, BMNH, ITNZ, JNNZ, LUNZ, MONZ, Decr. 1911. (hand-written) / New Zealand Broun Coll. NZAC, PHNZ). 1922-482 (typed) / Lecanomerus labralis. [male sym- Geographic distribution (Map p. 151). North Island: bol]. (hand-written).” Perfect condition. Synonymised AK, BP, CL, GB, HB, ND, WI, WN, WO. South Island: by Moore, in Moore et al., 1987: 227. DN, MC, NN, SC. Offshore Islands: CH. Adventive. Nemaglossa mastersi: Sloane, 1920: 137. Nemaglossa stenopus: Sloane, 1920: 137. Extralimital range: Australia (including Tasmania). First New Nemaglossa labralis: Sloane, 1920: 137. Zealand record: Howick, AK, 1880s (Broun, 1886: 880; as Acupalpus (Egadroma) vestigialis: Csiki, 1932: 1242. Lecanomerus stenopus). Well established. Lecanomerus vestigialis: Noonan, 1976: 7 and Moore, in Ecology. Lowland. Sand dunes, cultivated fields (maize, Moore et al., 1987: 227. lucerne, hay), pastures, and roadsides. Open ground; sandy Description. Body length: 4.0–4.3 mm. Strongly convex soil sparsely or moderately vegetated by grass, weeds, or (more so than verticalis). Black; appendages mostly red- herbs. Nocturnal; sheltering during the day in leaf litter and dish; antennomeres 3–11, apex of tibiae, and tarsi dark in burrows dug around the roots of plants (Muehlenbeckia, brown. Generally glabrous and smooth. Microsculpture Lupinus, Ammophila, Spinifex). Gregarious. Biology. isodiametric on head, moderately transverse on pronotum Seasonality: throughout the year. Tenerals: October–Janu- and elytra. Shiny, without metallic lustre; elytra slightly ary, March. Predacious (Moore et al., 1987: 228). Food, in iridescent. Head. Moderately large, narrower across eyes captivity: small scarabaeids and lygaeid nymphs. Occa- than pronotal apex; flat anteriorly, slightly convex sionally infested with mites and fungi (Laboulbeniales). posteriorly. Mandibles short, strongly curved forward. Dispersal power. Elytra free along suture. Macropterous. Labrum strongly transverse; apex slightly emarginate me- Occasional flier. Moderate runner. Good burrower. Col- dially. Eyes narrowly separated from buccal fissure lecting techniques. Pitfall trapping, raking leaf litter, and ventrally (by 0.5× maximum width of antennal scape). digging at the base of plants. Frons with clypeo-ocular prolongations complete. Anten- References. May, 1965: 202 (biology); Valentine, 1967: nae moderately long, reaching pronotal base; antennal scape 1102 (biology); Harris, 1970: 48, 53, 55 (distribution, ecol- about 2× longer than its maximum width; pubescence start- ogy); Cameron & Butcher, 1980: 115-116 (distribution, ing from basal 1/3 of antennomere 3. Mentum with a tooth ecology, biology); Moore et al., 1987: 227–228 (distribu- medially, moderately shorter than lateral lobes. Paraglossae tion, ecology, biology, dispersal power); Kuschel, 1990: as long as ligula. Palpi with last segment fusiform, with 24, 40 (distribution, ecology, biology, dispersal power); sparse, very short pubescence. Thorax. Pronotum (Fig. Townsend, 1994: 9, 11 and 1997: 17 (distribution, ecol- 154) very transverse, widest about middle (before middle ogy); Larochelle & Larivière, 2001: 121 (taxonomy, distri- in verticalis); sides converging toward base, not sinuate; bution, ecology, biology, dispersal power). base rather straight, moderately narrower than elytral base; apex concave; lateral depressions absent; anterior angles Remarks. See under L. verticalis. moderately developed, obtuse; posterior angles moderately developed, broadly rounded; basal foveae shallow, ill-de- fined; punctuation feebly developed basally. Metepisterna Genus Syllectus Bates, 1878 E longer than wide. Legs. Male mesotarsi dilated laterally Syllectus Bates 1878b: 191. Type species: Syllectus anomalus and spongily pubescent ventrally. Tarsi pubescent (with a Bates, 1878b, by monotypy. few setae) dorsally; metatarsomere 5 pubescent (with 6 Description. Body length: 4.7–8.0 mm. Forebody (head setae) ventrally; metatarsomere 1 as long as metatarsomeres and thorax) without sparse setiferous micropores dorsally. 2+3. Elytra. Widest about middle; slightly less elongate Forebody (head and thorax) much narrower than elytra Fauna of New Zealand 53 69

(contrary to other Harpalini genera, except Pholeodytes). Pelmatellina based on male protarsi that are spongily pu- Antennae and legs very long (contrary to other Harpalini bescent ventrally, the genus Syllectus superficially resem- genera, except Pholeodytes). Head. Mandibles very long bles the stenolophine genus Pholeodytes in having a (about 5x their maximum width), slightly curved forward, forebody much narrower than elytra, very long antennae acute apically. Labrum moderately transverse; apex straight and legs, and segment 4 of protarsi and mesotarsi with or slightly convex. Eyes moderately large and convex, reach- membranous laminae. These characters probably indicate ing eyes, or, strongly reduced and flat, consisting of oblit- an ecomorphological convergence in these cave-inhabiting erated facets and narrowly separated from buccal fissure taxa. ventrally (by about 0.5× maximum width of antennal scape). Tempora not inflated. Frons with or without clypeo-ocu- lar prolongations. Antennal pubescence starting from base Key to species of Syllectus of antennomere 3. Mentum with a tooth medially, as long 1 Eyes (Fig. 213) strongly reduced, flat, consisting of as lateral lobes (contrary to a longer medial tooth in obliterated facets. Pronotum (Fig. 157) elongate (about Pholeodytes). Mentum and submentum separated by com- 1.5× longer than wide). Frons without clypeo-ocular plete transverse suture. Paraglossae longer than ligula. Palpi prolongations. Interval 3 without setiferous puncture with last segment fusiform, not truncate apically, glabrous behind middle ...... (p. 71)... gouleti new species or with sparse, moderately long pubescence; penultimate — Eyes (Fig. 108) normally developed. Pronotum (Fig. segment of labial palpi bisetose on anterior margin. Tho- 155) quadrate (about as long as wide) or subrectangular rax. Pronotum quadrate (about as long as wide), (slightly longer than wide; Fig. 156). Frons with clypeo- subrectangular (slightly longer than wide) or elongate (about ocular prolongations (Fig. 108). Interval 3 with a 1.5x longer than wide; gouleti); base straight, much nar- setiferous puncture behind middle (Fig. 108)...... 2 rower than elytral base; lateral beads complete; anterior 2(1) Body length 6.0 mm or less. Eyes reaching buccal bead incomplete medially or absent (gouleti); posterior fissure ventrally (Fig. 21). Elytra oblong (Fig. 211). bead incomplete medially or absent (gouleti). Scutellum Last segment of palpi pubescent [Pronotum (Fig. 155)] visible. Apex of prosternal lobe glabrous. Legs. Metafemora ...... (p. 69)... anomalus Bates with 2–4 long setae on posterior margin. Male protarsi dilated laterally and spongily pubescent ventrally; mesotarsi — Body length 7.5 mm or more. Eyes not reaching buccal slightly dilated, but not spongily pubescent ventrally. Seg- fissure ventrally (Fig. 20). Elytra elliptical (Fig. 212). ment 4 of protarsi and mesotarsi of both sexes with 2 Last segment of palpi glabrous [Pronotum (Fig. 156)] membranous laminae (projecting laterally and anteriorly, ...... (p. 70)... magnus Britton as in Pholeodytes). Tarsi glabrous or with metatarsi par- tially pubescent dorsally; metatarsomere 5 glabrous or Syllectus anomalus Bates, 1878 E pubescent ventrally; metatarsomere 1 as long as Figures 72, 108, 155, 211; Map p. 151 metatarsomeres 2+3. Elytra. Interneurs complete, con- Syllectus anomalus Bates, 1878b: 192. Lectotype (here des- sisting of striae (contrary to interneurs incomplete, con- ignated): male (MNHN) labelled “Auckland N. Zeal. sisting of rows of punctures in Pholeodytes). Rows of (hand-written) / Syllectus anomalus Bates (hand-written) setiferous punctures absent on intervals 3, 5, and 7, and on / LECTOTYPE Syllectus anomalus Bates, 1878 desig- interneur 2. Umbilicate setiferous series of interval 9 sepa- nated by Larochelle & Larivière 2004 (red label; typed).” rated into two major groups, with posterior group further Good condition. Paralectotype: 1 female (MNHN) bear- divided into 2 subgroups. Abdomen. Ventrites 2+3 of male ing blue paralectotype label. without a setiferous fovea. Ventrites 5+6 of both sexes Description. Body length: 4.7–6.0 mm. Slightly convex. without short setae, with paired ambulatory setae only. Black; antennae, palpi, and legs brownish red. Generally Aedeagus. Lateral view: slightly arcuate or almost straight. glabrous and smooth. Microsculpture moderately trans- Dorsal view: symmetrical (with ostium dorsal, not de- verse, almost absent on head and pronotum, very trans- flected laterally); dorsal membranous area wide or narrow, verse (with microlines) on elytra. Shiny, without metallic extending to basal bulb or almost; apical disc absent. Inter- lustre; elytra iridescent. Head. Narrow, although as wide nal sac armed or unarmed. across eyes as pronotal apex; flat anteriorly, slightly con- vex posteriorly. Labrum with apex straight. Eyes moder- Geographic distribution. New Zealand (endemic). ately large, convex, reaching buccal fissure ventrally. Frons References. Hudson, 1934: 177 (list); Britton, 1964a: with clypeo-ocular prolongations complete. Antennae 629–631 (taxonomy); Noonan, 1976: 7 (taxonomy); moderarely long, reaching elytral base; antennal scape about Larochelle & Larivière, 2001: 121–122 (catalogue). 2× longer than its maximum width. Last segment of palpi Remarks. Although clearly a member of the subtribe with sparse, moderately long pubescence. Thorax. 70 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Pronotum (Fig. 155) quadrate, narrow, widest before mid- 2001: 121 (taxonomy, distribution, ecology, biology, dis- dle; sides converging toward base, slightly sinuate; base persal power). straight medially, angled forward laterally; apex concave; Remarks. Bates’ original description was apparently based lateral depressions widening posteriorly; anterior and pos- on specimens from both sexes (“[male symbol] [female terior beads incomplete medially; anterior angles feebly symbol]”) from “Auckland, New Zealand.” Two syntypes developed, subrectagular; posterior angles strongly devel- (1 male, 1 female) were obtained from the Bates collection oped, subrectangular; basal foveae deep, narrow; punctua- in Paris (MNHN); the male specimen which bears Bates’ tion feebly developed. Metepisterna longer than wide. Legs. determination label, is here selected as lectotype to pre- Metafemora with 2 long setae on posterior margin. Tarsi serve stability of nomenclature in the future. This species glabrous dorsally; metatarsomere 5 pubescent (with 2 se- is more widely distributed than the two other known tae) ventrally. Elytra. Oblong. Widest behind middle. Syllectus species. Syllectus anomalus can be found through- Shoulders well developed, rounded, without a tooth. Sub- out both main islands of New Zealand, mostly outside apical sinuations feeble. Sutural apices angulate. Scutellar caves, while the other species are restricted to caves in the striole absent. Interneurs moderately deep, finely punc- Northwest Nelson and Buller areas. tate. Intervals impunctate, slightly convex. Interval 3 with a setiferous puncture behind middle. Aedeagus (Fig. 72). Lateral view: slightly arcuate; apex broadly triangular; main Syllectus magnus Britton, 1964 E shaft narrower than in magnus. Dorsal view: symmetrical Figures 73, 156, 212; Map p. 151 (with ostium dorsal, not deflected laterally); dorsal mem- Syllectus magnus Britton, 1964a: 629. Holotype: male branous area very wide, extending almost to basal bulb. (NZAC) labelled “Type (circular red-bordered label; typed) Internal sac unarmed. / NEAR ENTRANCE ‘The Drain: ‘Profanity Cave’ Buller Material examined. 181 specimens, including types River near Inangahua 21.10.61. Coll. C. Coates. (hand- (AMNZ, CMNZ, ITNZ, JNNZ, LUNZ, MNHN, written) / HOLOTYPE [male symbol] Syllectus magnus mihi (hand-written) E.B. Britton det. 1963 (typed, ex- MONZ, NZAC, UCNZ). cept for number 3).” Perfect condition. There should be Geographic distribution (Map p. 151). North Island: one paratype in BMNH. AK, BP, CL, GB, HB, ND, RI, TK, TO, WA, WI, WN, Syllectus spelaeus Britton, 1964a: 631. Holotype: male WO. South Island: BR, DN, FD, KA, MB, MC, NC, NN, (NZAC) labelled “Type (circular red-bordered label; typed) SD, SL, WD. / Nile River Cave Charleston 27.10.63 P. R. Kettle (hand- written) / HOLOTYPE Syllectus spelaeus mihi (hand- Ecology. Lowland, montane, subalpine, alpine. Stream banks written) E. B. Britton det. 1964 (typed, except for number (brooks, rills, rivers) crossing cool wet forests (beech, 4).” Perfect condition. There are 3 paratypes in NZAC. broadleaf), tree plantations (pine), and scrublands; seep- New synonym. age edges, swamps, mud flats, screes, gardens; caves (occa- Description. Body length: 7.5–8.0 mm. Moderately con- sionally). Open or shaded ground; wet, clayey, muddy or vex. Brown; head and pronotum reddish brown; elytra gravelly soil, bare or sparsely vegetated with grass. Crep- entirely dark brown or black with reddish brown sides and uscular or nocturnal; sheltering during the day under small sutures; antennae, palpi, and legs pale yellowish brown. stones (mostly), clay clods, in heaps of dead leaves and Generally glabrous and smooth. Microsculpture well de- mud, and under logs. Gregarious. Biology. Seasonality: veloped, isodiametric on head, moderately transverse on September–April, July–August. Tenerals: February– pronotum, and very transverse (with microlines) on elytra. March. Predators: trout. Occasionally infested with mites. Shiny, without metallic lustre; elytra iridescent. Head. Dispersal power. Elytra free (macropterous) or fused Narrow, although as wide across eyes as pronotal apex; (brachypterous) along suture. Occasional flier (at dusk and flat anteriorly, slightly convex posteriorly. Labrum with to artifical lights at night). Moderate runner. Occasional apex slightly convex. Eyes moderately large, convex, nar- climber (on plants, shrubs, trees). Collecting techniques. rowly separated from buccal fissure ventrally (by about Turning stones, collecting at night with a torch, inspecting 0.5× maximum width of antennal scape). Frons with clypeo- heaps of dead leaves and mud, sweeping vegetation, light ocular prolongations complete. Antennae moderately long, trapping. reaching elytral base; antennal scape about 2× longer than References. May, 1962: 60 and 1972: 571 (distribution, its maximum width. Last segment of palpi glabrous. Tho- ecology, biology); May, 1963: 192 (ecology); Townsend, rax. Pronotum (Fig. 156) subrectangular (slightly longer 1974: 430 (ecology); Britton, 1964a: 631 (taxonomy); than wide), narrow, widest before middle; sides converging Johns, 1980: 63 (distribution, ecology); Kuschel, 1990: toward base, slightly sinuate; base straight medially, an- 24, 40 (distribution, ecology, biology, dispersal power); gled forward laterally; apex straight; lateral depressions Townsend, 1997: 17 (distribution); Larochelle & Larivière, widening posteriorly; anterior and posterior beads incom- Fauna of New Zealand 53 71 plete medially; anterior angles feebly developed, obtuse; brous and smooth. Microsculpture almost absent, posterior angles strongly developed, subrectangular; basal isodiametric on head, very transverse (with microlines) on foveae deep, ill-defined; punctuation feebly developed. pronotum and elytra. Shiny, without metallic lustre; Metepisterna longer than wide. Legs. Metafemora with 2 pronotum and elytra iridescent. Head. Narrow, although long setae on posterior margin. Pro- and mesotarsi gla- as wide across eyes as pronotal apex; flat anteriorly, slightly brous dorsally, metatarsi pubescent dorsally; metatarsomere convex posteriorly. Labrum with apex slightly convex me- 5 glabrous ventrally. Elytra. Elliptical. Widest behind mid- dially. Eyes strongly reduced, flat, consisting of obliter- dle. Shoulders feebly developed, rounded, without a tooth. ated facets, narrowly separated from buccal fissure Subapical sinuations feeble. Sutural apices rounded. ventrally (by about 0.5× maximum width of antennal scape). Scutellar striole absent. Interneurs moderately deep, al- Frons without clypeo-ocular prolongations. Antennae very most impunctate. Intervals impunctate, slightly convex. long, reaching middle of elytra; antennal scape elongate, Interval 3 with a setiferous puncture behind middle. about 3× longer than its maximum width. Last segment of Aedeagus (Fig. 73). Lateral view: slightly arcuate; apex palpi glabrous. Thorax. Pronotum (Fig. 157) elongate broadly triangular; wider than in anomalus. Dorsal view: (about 1.5× longer than wide), widest about middle; sides symmetrical (with ostium dorsal, not deflected laterally); converging toward base, slightly sinuate; base straight dorsal membranous area very wide, extending almost to throughout; apex straight; lateral depressions widening basal bulb, with a medial swelling subapically. Internal sac posteriorly; anterior and posterior beads absent; anterior armed. angles feebly developed, rounded; posterior angles strongly Material examined. 51 specimens, including types developed, subrectangular; basal foveae deep, wide; punc- (BMNH, ITNZ, LUNZ, NZAC, UCNZ). tuation feebly developed. Metepisterna longer than wide. Legs. Metafemora with 3–4 long setae on posterior mar- Geographic distribution (Map p. 151). South Island: gin. Tarsi glabrous dorsally; metatarsomere 5 glabrous BR, NN. ventrally. Elytra. Elliptical. Widest behind middle. Shoul- Ecology. Lowland. Cave entrances: in the twilight zone ders feebly developed, rounded, without a tooth. Subapi- (troglophilous species). In flood debris. Biology. cal sinuations feeble. Sutural apices angulate. Scutellar striole Seasonality: September–June. Dispersal power. Elytra absent. Interneurs moderately deep, impunctate. Intervals fused along suture. Subapterous. Fast runner. Collecting impunctate, slightly convex. Interval 3 without setiferous techniques. Using a head lamp or torch. puncture behind middle. Aedeagus (Fig. 74). Atypical for References. Britton, 1964a: 631 (including spelaeus; dis- genus. Dorsal view: symmetrical (with ostium dorsal, not tribution, ecology); Townsend, 1974: 430 (as spelaeus; deflected laterally); dorsal membranous area narrow, ex- ecology) and 1997: 17–18 (including spelaeus; distribu- tending to basal bulb. Internal sac armed. tion, ecology; Larochelle & Larivière, 2001: 121–122 (in- Material examined. 17 specimens, including types (ITNZ, cluding spelaeus; taxonomy, distribution, ecology, biology, LUNZ, NZAC). dispersal power). Geographic distribution (Map p. 151). South Island: BR– Remarks. Syllectus spelaeus is synonymised with S. Bamboo Cave, Tiropahi Valley. Dry Valley Cave, Bullock magnus on the basis of the male genitalia. Syllectus magnus Creek, Punakaiki. Fox River Cave. Metro Cave, Charleston. is a taxon that varies somewhat morphologically (espe- Nile River Cave, Charleston. Tetahi Cave, Charleston. Xanadu cially the pronotum and body colour) within and between Cave, Bullock Creek, Paparoa National Park. populations across its range. Ecology. Lowland. Caves (troglobitic species). Running on mud banks. Biology. Seasonality: January–February, Syllectus gouleti new species E April, July. Tenerals: December. Dispersal power. Elytra fused along suture. Subapterous. Fast runner. Collecting Figures 74, 157, 213; Map p. 151 Syllectus gouleti Larochelle & Larivière, new species. techniques. Using a head lamp or torch. Holotype: male (NZAC) labelled “Metro Cave June J.I. Remarks. Although this species superficially resembles Townsend [hand-written] / JI Townsend Collection members of the genus Pholeodytes, it bears the characteris- (typed)/ HOLOTYPE [male symbol] Syllectus gouleti Larochelle & Larivière, 2004 (red label; typed).” tic features of Syllectus (principally the ventral pubes- Paratypes: 1 male (NZAC) and 1 female (LUNZ) from cence of male tarsi). The aedeagus is however unusual among the same locality as the holotype, bearing blue paratype Syllectus. This species is named after our close friend and labels. colleague Henri Goulet (Agriculture and Agri-Food Canada, Description. Body length: 6.5–7.5 mm. Slightly convex. Ottawa, Canada) for his special help and encouragement in Depigmented (appearing pale in colour). Generally gla- our entomological studies. 72 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Subtribe STENOLOPHINA 3(1) Elytral base and side with interneurs incomplete Diagnosis (New Zealand). Body length: 3.0–8.3 mm. Frons basally and laterally (Fig. 111). Clypeo-ocular usually with clypeo-ocular prolongations, seldom without prolongations incomplete toward eyes (Fig. 111). (Pholeodytes). Mentum usually with a tooth medially, sel- Pronotum (Fig. 163) suborbicular. Apex of prosternal dom without (Egadroma). Mentum and submentum usu- lobe glabrous (Fig. 2) ...(p. 77)... Haplanister Moore ally separated by complete transverse suture, seldom by — Elytral base and side with interneurs complete (Fig. laterally incomplete transverse suture (Euthenarus). Pe- 109–110). Clypeo-ocular prolongations complete (Fig. nultimate segment of labial palpi bisetose (with 2 setae). 110). Pronotum transverse, not suborbicular. Apex of Apex of prosternal lobe glabrous or pubescent (Egadroma, prosternal lobe pubescent ...... 4 Euthenarus). Male protarsi dilated laterally and biseriately 4(3) Ventrites 5+6 with numerous short setae, in addition pubescent ventrally, seldom unmodified (Haplanister). to paired ambulatory setae (Fig. 28). Mentum tooth Male mesotarsi dilated laterally and biseriately pubescent present (Fig. 14). Ventrites 2+3 of male with a setiferous ventrally (except 2 adventive Euthenarus and Haplanister). fovea (Fig. 28) ...... (p. 73)... Euthenarus Bates Metatarsomere 1 usually as long as metatarsomeres 2+3, — Ventrites 5+6 with paired ambulatory setae only (Fig. rarely shorter (Haplanister, some Euthenarus species). 27). Mentum tooth absent (Fig. 18). Ventrites 2+3 of Umbilicate setiferous series of interval 9 separated into 2 male without a setiferous fovea (Fig. 27) ...... major groups with posterior group divided further into 2 ...... (p. 72)... Egadroma Motschulsky subgroups or continuous. Aedeagus arcuate, usually sym- metrical (with ostium dorsal, not deflected laterally), sel- dom asymmetrical (with ostium slightly deflected to the Genus Egadroma Motschulsky, 1855 A left; Egadroma). Egadroma Motschulsky, 1855: 43. Type species: Carabus smaragdulus Fabricius, 1798, by monotypy. Geographic distribution. Worldwide. Stenolophus (Egadroma): Ganglbauer, 1892: 370. References. Noonan, 1976: 15–28 (taxonomy); Larochelle Acupalpus (Egadroma): Csiki, 1932: 1239. & Larivière, 2001: 126–128 (catalogue). Stenolophus (Egadroma): Noonan, 1976: 18. Egadroma: Serrano et al., 1994: 56. Remarks. The dilatation and pubescence of the male pro- and mesotarsi could not be studied for the genus Description (New Zealand). Body length: 5.0–6.5 mm. Kiwiharpalus which is only known from females. Forebody (head and thorax) without sparse setiferous micropores dorsally. Head. Mandibles short, strongly curved forward, blunt apically. Labrum strongly transverse; apex straight or slightly emarginate medially. Eyes moder- Key to genera of New Zealand Stenolophina ately large, convex, reaching buccal fissure ventrally. 1 Eyes (Fig. 112–113) strongly reduced, flat, consisting of Tempora not inflated. Frons with clypeo-ocular obliterated facets. Mandibles very long (5–6× their prolongations complete. Antennal pubescence starting from maximum width; Fig.113). Body depigmented middle of antennomere 2. Mentum without tooth medi- (appearing pale in colour; Fig. 220–225) ...... 2 ally. Mentum and submentum separated by complete trans- verse suture. Paraglossae longer than ligula. Palpi with last — Eyes (Fig. 109) normally developed. Mandibles shorter segment fusiform, not truncate apically, almost glabrous; (Fig. 109). Body pigmented (appearing dark in colour) penultimate segment of labial palpi bisetose on anterior ...... 3 margin. Thorax. Pronotum transverse; base convex, mod- 2(1) Segment 4 of protarsi and mesotarsi with 2 erately narrower than elytral base; lateral beads complete; membranous laminae (Fig. 25). Elytral interneurs anterior bead incomplete medially; posterior bead absent. incomplete, consisting of rows of punctures (Fig. 113). Scutellum visible. Apex of prosternal lobe pubescent. Legs. Forebody (head and thorax) much narrower than elytra Metafemora with 2 long setae on posterior margin. Male (Fig. 113). Body length 6.0 mm or more [Cave beetles. protarsi and mesotarsi dilated laterally and biseriately pu- South Island: NN] ...... (p. 80)... Pholeodytes Britton bescent ventrally. Segment 4 of protarsi and mesotarsi of both sexes without membranous laminae. Tarsi glabrous — Segment 4 of protarsi and mesotarsi without dorsally; metatotarsomere 5 pubescent ventrally (with 2 membranous laminae (Fig. 26). Elytral interneurs setae); metatarsomere 1 as long as metatarsomeres 2+3. complete, consisting of striae (Fig. 112). Forebody Elytra. Interneurs complete. Rows of setiferous punc- (head and thorax) at most moderately narrower than tures absent on intervals 3, 5, and 7, and on interneur 2. elytra (Fig. 112). Body length 3.5mm or less [Offshore Umbilicate setiferous series of interval 9 separated into 2 Islands: TH] ...... (p. 79)... Kiwiharpalus new genus major groups, with posterior group further divided into 2 Fauna of New Zealand 53 73 subgroups. Abdomen. Ventrites 2+3 of male without a elytral base; antennal scape about 3× longer than its maxi- setiferous fovea. Ventrites 5+6 of both sexes without short mum width. Thorax. Pronotum (Fig. 158) very transverse, setae, with paired ambulatory setae only. Aedeagus (Fig. widest before middle; sides converging toward base, not 75). Lateral view: slightly arcuate. Dorsal view: asym- sinuate; apex concave; lateral depressions raised, moder- metrical (with ostium deflected to the left); dorsal mem- ately large, slightly widening posteriorly; anterior angles branous area wide, extending almost to basal bulb; apical moderately developed, rounded; posterior angles feebly disc present. Internal sac armed. developed, broadly rounded; basal foveae shallow, wide, Geographic distribution. Ethiopian, Australian, Palearctic, extending to lateral beads; punctuation strongly developed and Oriental Regions, the Pacific Islands; New Zealand basally (particularly in basal foveae). Apex of prosternal (adventive). lobe with 5–6 long setae. Metepisterna longer than wide. Elytra. Widest behind midddle. Shoulders strongly devel- References. Noonan, 1976: 18 (taxonomy); Serrano et al., oped, rounded, without a tooth. Subapical sinuations fee- 1994: 56 (karyotype, taxonomy); Serrano & Galián, 1998: ble. Sutural apices angulate. Scutellar striole present. 198 (karyotype, taxonomy); Larochelle & Larivière, 2001: Interneurs shallow, deepening apically, impunctate. Inter- 126 (catalogue). vals impunctate, flat, becoming convex apically. Interval 3 Remarks. Serrano et al., (1994: 56) and Serrano & Galián with a setiferous puncture behind middle. Aedeagus (Fig. (1998: 198) separated Egadroma from Stenolophus Dejean 75). As for genus. on the basis of chromosome number, meiotic behaviour of Material examined. 23 non-type specimens (AMNZ, chromosomes, and geographic distribution, the latter taxon JNNZ, LUNZ, NZAC). being restricted to the Holarctic Region. The genus Egadroma is in need of revision. Geographic distribution (Map p. 151). North Island: AK, CL, ND. Adventive. Extralimital range: New Caledo- nia, Pacific Islands, Australia (including Tasmania). First Egadroma picea (Guérin-Méneville, 1830) A New Zealand records: Lake Ohia, ND, 1917 (NZAC) ; Figures 75, 109, 158, 214; Map p. 151 Redhill and Woodhill, AK, Rangiputa and Ruakaka, ND Acupalpus piceus Guérin-Méneville, 1830: Plate 1, Figure (Cameron & Butcher, 1980: 115–116). Well established. 12. Type locality: Port Jakson [=Port Jackson], New South Wales, Australia. Ecology. Lowland. Sandy pastures and sand dunes. Open Harpalus dingo Laporte de Castelnau, 1867: 111 (redescribed ground; dry, sandy, bare or sparsely vegetated soil. Noc- in 1868: 197). Type locality: Rockhampton, Queens- turnal; sheltering during the day in soil burrows dug at the land, Australia. Synonymised by Moore, in Moore et al., base of plants and under logs. Biology. Seasonality: Sep- 1987: 242. tember–March, August. Omnivorous, probably granivorous Homalosoma dingo: Gemminger & Harold, 1868: 329. (Moore et al., 1987: 242). Food, in captivity: Small Stenolophus politus Macleay, 1871: 103. Type locality: scarabaeids. Dispersal power. Elytra free along suture. Gayndah, Queensland, Australia. Synonymised by Moore, Macropterous. Occasional flier. Moderate runner. Good in Moore et al., 1987: 242. Stenolophus dingo: Chaudoir, 1878: 514. burrower. Effective as a colonist. Favoured by human ac- Stenolophus sexualis Fauvel, 1882: 270. Type locality: Ile tivities. Collecting techniques. Digging at the base of des Pins and Nouméa, New Caledonia. Synonymised by plants, turning logs and seashore debris, light trapping. Sloane, 1920b: 323. References. Moore et al., 1987: 242 (distribution, ecol- Stenolophus piceus: Cameron & Butcher, 1980: 115. ogy, biology, dispersal power); Cameron & Butcher, 1980: Stenolophus (Egadroma) piceus: Moore et al., 1987: 242. 115–116 (as Stenolophus piceus, distribution, ecology, bi- Egadroma piceus [sic]: Serrano et al., 1994: 56. ology); Larochelle & Larivière, 2001: 126 (taxonomy, dis- Description. Body length: 5.0–6.5 mm. Slightly convex. tribution, ecology, biology, dispersal power). Dark brown; pronotum with wide reddish or yellowish margins; elytra with a reddish spot (more or less distinct) on the shoulder; suture with posterior half yellowish red; Genus Euthenarus Bates, 1874 N antennomeres 2–3, palpi, and legs yellow. Generally gla- Euthenarus Bates 1874: 272 (redescribed in 1875: 310). Type brous and smooth. Microsculpture isodiametric on head, species: Euthenarus brevicollis Bates, 1874, designated moderately transverse on pronotum, very transverse (with by Noonan, 1976: 27. microlines) on male elytra, granulate on female elytra. Shiny, Euthenaris: Csiki, 1932: 1268 (incorrect subsequent spell- without metallic lustre; elytra iridescent in males, dull in ing). females. Head. Moderately large, narrower across eyes Description. Body length: 3.8–6.0 mm. Forebody (head than pronotal apex; flat anteriorly, slightly convex and thorax) without sparse setiferous micropores dorsally. posteriorly. Antennae moderately long, reaching about Head. Mandibles short, slightly curved forward, blunt 74 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) apically. Labrum strongly transverse; slightly emarginate Key to species of Euthenarus medially. Eyes moderately large, convex, reaching or al- 1 Eyes reaching buccal fissure ventrally (Fig. 21). Femora most reaching buccal fissure ventrally. Tempora not in- pale. Male mesotarsi unmodified ...... 2 flated. Frons with clypeo-ocular prolongations complete. Antennal pubescence starting from middle of antennomere — Eyes narrowly separated from buccal fissure ventrally × 2. Mentum with a tooth medially, moderately shorter than (by 0.7 maximum width of antennal scape; Fig. 20). lateral lobes. Mentum and submentum separated by later- Femora dark. Male mesotarsi dilated laterally (Fig. 215) ally incomplete transverse suture. Paraglossae longer than and biseriately pubescent ventrally (Fig. 13) ...... 3 ligula. Palpi with last segment fusiform, not truncate 2(1) Body length 4.0 mm or less. Pronotum (Fig. 161): apically, subglabrous; penultimate segment of labial palpi reddish brown, contrasting with dark head and elytra; bisetose on anterior margin. Thorax. Pronotum transverse; posterior angles rounded; base convex [Aedeagus (Fig. base convex or straight, moderately narrower than elytral 78)] ...... (p. 76)... bicolor Moore base; lateral beads complete; anterior and posterior beads — Body length over 4.0 mm. Pronotum (Fig. 162): piceous incomplete medially. Scutellum visible. Apex of prosternal brown with paler borders, not contrasting with colour lobe pubescent. Legs. Metafemora with 2 long setae on of head and elytra; posterior angles rectangular; base posterior margin. Male protarsi dilated laterally and straight [Aedeagus (Fig. 79)] ...... biseriately pubescent ventrally; male mesotarsi dilated lat- ...... (p. 77)... promptus (Erichson) erally and biseriately pubescent ventrally (brevicollis, 3(1) Subapical sinuations of elytra feeble (Fig. 29, puncticollis) or unmodified (bicolor, promptus). Segment 4 laterodorsal view). Pronotum (Fig. 159): posterior of protarsi and mesotarsi of both sexes without membra- angles obtuse-rounded; base strongly convex. Elytral nous laminae. Tarsi glabrous dorsally; metatarsomere 5 interneurs shallow, almost erased apically [Aedeagus glabrous ventrally; metatarsomere 1 shorter than or as long (Fig. 76)] ...... (p. 74)...brevicollis Bates as (promptus) metatarsomeres 2+3. Elytra. Interneurs com- plete. Rows of setiferous punctures absent on intervals 3, — Subapical sinuations of elytra strong (Fig. 30, 5, and 7, and on interneur 2. Umbilicate setiferous series of laterodorsal view). Pronotum (Fig. 160): posterior interval 9 separated into 2 major groups, with posterior angles almost rectangular; base slightly convex. Elytral group continuous. Abdomen. Ventrites 2+3 of male with a interneurs deeper, well impressed apically (Fig. 110) setiferous fovea medially (contrary to other Harpalini gen- [Aedeagus (Fig. 77)] ...... (p. 75)... puncticollis Bates era). Ventrites 5+6 of both sexes with numerous short setae, in addition to paired ambulatory setae (contrary to E other Harpalini genera). Aedeagus. Lateral view: strongly Euthenarus brevicollis Bates, 1874 or moderately arcuate. Dorsal view: symmetrical (with Figures 76, 159, 215; Map p. 152 ostium dorsal, not deflected laterally); dorsal membranous Euthenarus brevicollis Bates, 1874: 273 (redescribed in 1875: 311). Lectotype (here designated): male (MNHN) la- area wide or narrow (promptus), extending almost to basal belled “Canterby N. Zeal (hand-written) / Euthenarus bulb; apical disc present or absent. Internal sac armed or brevicollis Bates (hand-written) / LECTOTYPE unarmed. Euthenarus brevicollis Bates, 1878 designated by Geographic distribution. Australia (including Tasmania Larochelle & Larivière 2004 (red label; typed).” Good and Norfolk Island), New Zealand. condition. Euthenaris brevicollis: Csiki, 1932: 1268 (incorrect subse- References. Bates, 1874: 273 (taxonomy); Hudson, 1934: quent spelling). 177 (list); Noonan, 1976: 27–28 (taxonomy); Moore, 1985: Description. Body length: 5.0–6.0 mm. Moderately con- 252–253 (taxonomy, distribution); Townsend, 1997: 17 vex. Head, thorax, and femora piceous black; elytra dark (species diagnosis); Larochelle & Larivière, 2001: 126–127 brown; pronotal margins, elytral suture, apical 1/3 of inter- (catalogue). val 9, antennal base, and base and apex of palpi rufous. Remarks. This genus is in need of revision. Generally glabrous and smooth. Microsculpture isodiametric on head and pronotum, slightly transverse on elytra. Shiny, with aeneous or coppery lustre; elytra iri- descent in males, dull in females. Head. Moderately large, narrower across eyes than pronotal apex; flat anteriorly, slightly convex posteriorly. Eyes narrowly separated from buccal fissure ventrally (by about 0.7× maximum of antennal scape). Antennae moderately long, reaching elytral base; antennal scape about 2× longer than its maximum width. Fauna of New Zealand 53 75

Thorax. Pronotum (Fig. 159) moderately transverse, Bates and a locality label reading “Canterby [=Canter- subrectangular, widest before middle; sides converging to- bury] N. Zeal”. The other MNHN specimens were col- ward base, not sinuate; base strongly convex (more so than lected in “Christchurch” and do not bear a Bates’ determi- in puncticollis); apex straight; lateral depressions absent; nation label. In addition, two male specimens collected in anterior angles feebly developed, rounded; posterior an- Oakden’s (near Lake Coleridge) in 1873 were located in the gles moderately developed, obtuse-rounded; basal foveae Wakefield material deposited in the Canterbury Museum moderately deep, wide; punctuation strongly developed (CMNZ). The locality labels for the Canterbury specimen (in basal foveae). Apex of prosternal lobe with 5–6 long (MNHN) and Oakden’s specimens (CMNZ) have been setae. Metepisterna longer than wide. Legs. Male mesotarsi written by different hands, which casts doubts about dilated laterally and biseriately pubescent ventrally. whether Wakefield was also the collector of the Canter- Metatarsomere 1 shorter than metatarsomeres 2+3. Elytra. bury specimen (MNHN). On the other hand, it is not clear Widest about middle. Shoulders strongly developed, either that the Oakden’s specimens collected by Wakefield rounded, without a tooth. Subapical sinuations feeble. (CMNZ) were part of the syntype series, i.e., they may or Sutural apices angulate. Scutellar striole absent. Interneurs may not have been seen by Bates. Because the Canterbury complete, shallow, almost erased apically, impunctate. In- specimen (MNHN) is more likely to have been part of tervals impunctate, flat. Interval 3 with a setiferous punc- Bates’ original type series — specimens were not always ture behind middle. Aedeagus (Fig. 76). Lateral view: labelled with precise localities in those days — it is here strongly arcuate; apex narrowly pointed; apical 1/2 of main designated as lectotype to preserve stability of nomencla- shaft narrowly triangular. Dorsal view: symmetrical (with ture in the future. This species is morphologically close to ostium dorsal, not deflected laterally); dorsal membranous E. puncticollis, the only other endemic Euthenarus. area very wide, extending almost to basal bulb; apical disc present, rounded at tip, less than 0.5× as long as wide. E Internal sac armed. Euthenarus puncticollis Bates, 1874 Material examined. 79 specimens, including type Figures 77, 110, 160, 216; Map p. 152 Euthenarus puncticollis Bates, 1874: 273 (redescribed in 1875: (AMNZ, BBNZ, CMNZ, JNNZ, LUNZ, MNHN, 311). Lectotype (here designated): male (MNHN) la- MONZ, NZAC, OMNZ, PHNZ, UCNZ). belled “Auckland N. Zeald (hand-written) / Euthenarus Geographic distribution (Map p. 152). South Island: CO, puncticollis Bates (hand-written) / LECTOTYPE DN, FD, KA, MC, MK, NC, OL, SC, SL. Stewart Island. Euthenarus puncticollis Bates, 1878 designated by Larochelle & Larivière 2004 (red label; typed).” Good Ecology. Lowland, montane, subalpine. Borders of lakes, condition. Paralectotype: 1 female (MNHN) bearing blue ponds, and slowly running streams; dried-up stream beds, paralectotype label. swamps; wet pastures and cultivated fields. Open ground; Euthenaris puncticollis: Csiki, 1932: 1268 (incorrect subse- wet, muddy, bare or vegetated soil. Nocturnal; sheltering quent spelling). during the day under logs and stones. Gregarious. Biology. Description. Body length: 5.0–6.0 mm. Moderately con- Seasonality: September–March, June. Tenerals: late Janu- vex. Head, thorax, and femora piceous black; elytra dark ary (MK). Occasionally infested with mites. Dispersal brown; pronotal margins, elytral suture, apical 1/3 of inter- power. Elytra free along suture. Macropterous. Occasional val 9, antennal base, and base and apex of palpi rufous. in seashore drift material, which indicates flight ability. Generally glabrous and smooth. Microsculpture Moderate runner. Good burrower. Occasional climber (on isodiametric on head and thorax, slightly transverse on grass, sedges, currants). Collecting techniques. Pitfall elytra. Shiny, with aeneous or coppery lustre; elytra iri- trapping, turning logs and stones, sweeping plants. descent in males, dull in females. Head. Moderately large, References. Johns, 1986: 31 (distribution, ecology); narrower across eyes than pronotal apex; flat anteriorly, Townsend, 1997: 17 (taxonomy, distribution); Emberson, slightly convex posteriorly. Eyes narrowly separated from 1998: 30 (taxonomy, distribution, ecology, biology); buccal fissure ventrally (by about 0.7× maximum width of Larochelle & Larivière, 2001: 127 (taxonomy, distribution, antennal scape). Antennae moderately long, reaching elytral ecology, biology, dispersal power). base; antennal scape about 2× longer than its maximum Remarks. Bates’ original description was based on an width. Thorax. Pronotum (Fig. 160) moderately trans- unspecified number of specimens of both sexes (“[male verse, subrectangular, widest before middle; sides converg- symbol, female symbol]”) from “Lake Coleridge” (situ- ing toward base, slightly sinuate; base slightly convex (less ated in the Alps in Canterbury), and collected by C. M. so than in brevicollis); apex straight; lateral depressions Wakefield. Seven specimens of E. brevicollis were obtained absent; anterior angles feebly developed, rounded; poste- from the Bates collection in Paris (MNHN), one of which rior angles strongly developed, almost rectangular; basal (a male specimen) bears a determination label written by foveae very deep, wide; punctuation strongly developed 76 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

(in basal foveae). Apex of prosternal lobe with 5–6 long Euthenarus bicolor Moore, 1985 A first New Zealand setae. Metepisterna longer than wide. Legs. Male mesotarsi record dilated laterally and biseriately pubescent ventrally. Figures 78, 161, 217; Map p. 152 Metatarsomere 1 shorter than metatarsomeres 2+3. Elytra. Euthenarus bicolor Moore, 1985: 252. Type locality: Black Widest about middle. Shoulders strongly developed, Mt., A.C.T., Australia. rounded, with a tooth. Subapical sinuations strong. Sutural Description. Body length: 3.8–4.0 mm. Moderately con- apices angulate. Scutellar striole absent. Interneurs shal- vex. Head black; pronotum reddish brown; elytra dark low, moderately deep apically, impunctate. Intervals brown; legs, antennal base, and palpi yellowish. Generally impunctate, slightly convex. Interval 3 with a setiferous glabrous and smooth. Microsculpture vestigial on head puncture behind middle. Aedeagus (Fig. 77). Lateral view: and pronotum, very transverse (with microlines) on elytra. strongly arcuate; apex narrowly pointed (more attenuate Shiny, without metallic lustre; elytra iridescent. Head. than in brevicollis); apical half of main shaft not triangular. Moderately large, narrower across eyes than pronotal apex; Dorsal view: symmetrical (with ostium dorsal, not de- flat anteriorly, slightly convex posteriorly. Eyes reaching flected laterally); dorsal membranous area wide, extending buccal fissure ventrally. Antennae moderately long, reach- almost to basal bulb; apical disc present, rounded at tip, ing about elytral base; antennal scape about 2×longer than more than 0.5× as long as wide. Internal sac armed. its maximum width. Thorax. Pronotum (Fig. 161) moder- Material examined. 250 specimens, including types ately transverse, widest before middle; sides converging (AMNZ, CMNZ, ITNZ, JNNZ, LUNZ, MNHN, toward base, not sinuate; base slightly convex; apex straight; MONZ, NZAC, UCNZ). lateral depressions absent; anterior angles feebly devel- Geographic distribution (Map p. 152). North Island: oped, rounded; posterior angles feebly developed, rounded; AK, CL, GB, HB, ND, RI, TK, TO, WA, WI, WN, WO. basal foveae shallow, wide; punctuation strongly devel- South Island: BR, NN, SD, WD. Offshore Islands: CH. oped (in basal foveae). Apex of prosternal lobe with 5–6 long setae. Metepisterna longer than wide. Legs. Male Ecology. Lowland, montane, subalpine. Borders of lakes, mesotarsi unmodified. Metatarsomere 1 shorter than ponds, flax swamps, slowly running rivers and brooks; metatarsomeres 2+3. Elytra. Widest about middle. Shoul- mud flats and wet pastures. Open ground; wet, muddy ders strongly developed, rounded, without a tooth. Sub- (mostly) or sandy soil with sparse vegetation (e.g., Juncus). apical sinuations feeble. Sutural apices angulate. Scutellar Nocturnal; sheltering during the day in burrows at the base striole absent. Interneurs deep, impunctate. Intervals of plants and under embedded branches and logs (mostly); impunctate, flat. Interval 3 with a setiferous puncture be- in soil cracks, leaf litter, and moss. Gregarious. Biology. hind middle. Aedeagus (Fig. 78). Lateral view: moderately Seasonality: throughout the year. Tenerals: October arcuate; apex triangular; apical half of main shaft broadly (rarely), February–April, June. Occasionally infested with triangular. Dorsal view: symmetrical (with ostium dorsal, mites and fungi (Laboulbeniales). Dispersal power. Elytra not deflected laterally); dorsal membranous area very wide, free along suture. Macropterous. Regular in seashore drift extending almost to basal bulb; apical disc absent. Internal material, which indicates flight ability. Moderate runner. sac unarmed. Much smaller and stouter than in other Good burrower. Occasional climber (on shrubs and trees). Collecting techniques. Digging at base of Juncus-tufts, Euthenarus species. turning embedded branches and logs, pouring water over Material examined. 144 non-type specimens (AMNZ, ground, using pan traps, turning seashore drift material. LUNZ, NZAC). References. Barratt & Patrick, 1987: 82 (distribution, ecol- Geographic distribution (Map p. 152). North Island: ogy); Townsend, 1994: 9, 11–13 (distribution, ecology); AK, BP, CL, GB, HB, ND, TO, WO. South Island: NN. Townsend, 1997: 17 (taxonomy, distribution); Larochelle Adventive. Extralimital distribution: Australia (including & Larivière, 2001: 127 (taxonomy, distribution, ecology, Tasmania and Norfolk Island). First New Zealand record: biology, dispersal power). Auckland, AK, 1945 (NZAC). Well established. Remarks. Bates’ original description was based on speci- Ecology. Lowland. Borders of eutrophic marshes and mens from both sexes “[male symbol, female symbol]”, ponds; swamp forests; borders of slowly running rivers including “several examples” from “Auckland”. Six speci- and brooks; mud flats, roadside ditches, and wet vacant mens collected in Auckland were obtained from the Bates lots. Open ground; soft, wet, muddy (mostly) or sandy collection in Paris (MNHN), two of which (a male and a soil with sparse or moderate vegetation (mostly Juncus, female) bear a determination label written by Bates; these also Carex, Typha, or weeds). Mostly nocturnal; occasion- two specimens are here considered syntypic and labelled ally running during the day in the spring sunshine; usually as lectotype and paralectotype to preserve stability of sheltering during the day in burrows at the base of plants nomenclature in the future. See under E. brevicollis. (mostly), in plant debris and leaf litter, under pieces of Fauna of New Zealand 53 77 wood and stones. Gregarious. Biology. Seasonality: Sep- Widest about middle. Shoulders strongly developed, tember–April, August. Omnivorous (Moore et al., 1987). rounded, without a tooth. Subapical sinuations feeble. Occasionally infested with mites. Dispersal power. Elytra Sutural apices angulate. Scutellar striole absent. Interneurs free along suture. Macropterous. Regular flier (to artificial shallow, well impressed apically, impunctate. Intervals lights at night). Regular in seashore drift material, which impunctate, flat. Interval 3 with a setiferous puncture be- also indicates flight ability. Moderate runner. Good bur- hind middle. Aedeagus (Fig. 79). Lateral view: moderately rower. Occasional climber (on rushes and trees). Clearly arcuate; apex narrowly truncate; apical 1/2 of main shaft effective as a colonist. Favoured by human activities. Col- not triangular, inflated ventrally. Dorsal view: symmetrical lecting techniques. Pouring water over ground, treading (with ostium dorsal, not deflected laterally); dorsal mem- soil with feet, treading vegetation into water, light trap- branous area narrow, extending almost to basal bulb; apical ping, sweeping vegetation, turning seashore drift material. disc present, triangular. Internal sac armed. References. Moore, 1985: 253 (taxonomy, distribution, Material examined. 113 non-type specimens (ITNZ, dispersal power). Moore et al., 1987: 244 (synonymy, JNNZ, LUNZ, MONZ, NZAC). distribution, ecology, biology, dispersal power). Geographic distribution (Map. 152). North Island: HB, Remarks. The reddish brown pronotum contrasting with TO, WA, WI, WN. South Island: BR, KA, MB, MC, NN, the dark head and elytra, and the body shape make this SD. Adventive. Extralimital distribution: Australia (includ- species superficially similar to Lecanomerus atriceps. Both ing Tasmania and Norfolk Island). First New Zealand species have often been confused in the past, but they are record: The Glen, Nelson, NN, 1950 (NZAC). Well estab- easily distinguished based on male tarsal characters lished. (biseriately pubescent ventrally in E. bicolor, spongily Ecology. Lowland. Stream banks, swamps, roadside banks, pubescent in L. atriceps) and ventral pubescence (ventrites and wet pastures. Open ground; wet, sandy soil with sparse 5+6 with numerous setae and male setiferous fovea of vegetation (grass, sedges). Nocturnal; sheltering during the ventrites 2+3 present in E. bicolor, such pubescence lack- day in burrows at the base of plants (mostly), under plants ing in L. atriceps). and logs. According to Moore (1985), this species “is gen- erally associated with wet ground, under or among subaquatic plants.” Gregarious. Biology. Seasonality: Sep- Euthenarus promptus (Erichson, 1842) A first New tember–March, July–August. Tenerals: March (abundant), Zealand record July. Omnivorous (Moore et al., 1987). Dispersal power. Figures 79, 162, 218; Map p. 152 Elytra free along suture. Macropterous. Regular flier (to Harpalus promptus Erichson, 1842: 126. Type locality: Tas- artificial lights at night). Regular in seashore drift material, mania, Australia. which also indicates flight ability. Moderate runner. Good Euthenarus promptus: Sloane, 1920: 138. burrower. Occasional climber (on rushes and grass). Clearly Description. Body length: 4.0–6.0 mm. Moderately con- effective as a colonist. Favoured by human activities. Col- vex. Head, pronotum, and elytra piceous brown; pronotal lecting techniques. Pouring water over ground, treading borders, elytral suture, apical 1/3 of interval 9, pronotal soil with feet, treading vegetation into water, uprooting and elytral epipleura, antennal base, palpi, and legs reddish plants, light trapping, sweeping vegetation, turning sea- yellow. Generally glabrous and smooth. Microsculpture shore drift material. strongly transverse dorsally. Shiny, with green metallic References. Moore, 1985: 252–253 (taxonomy, distribu- lustre. Head. Moderately large, narrower across eyes than tion, ecology, dispersal power). Moore et al., 1987: 245 pronotal apex; flat anteriorly, slightly convex posteriorly. (synonymy, distribution, ecology, biology, dispersal Eyes reaching buccal fissure ventrally. Antennae moder- power). ately long, reaching elytral base; antennal scape about 2× longer than its maximum width. Thorax. Pronotum (Fig. 162) moderately transverse, subrectangular, widest before Genus Haplanister Moore, 1996 A middle; sides converging toward base, slightly sinuate; base Haplanister Moore, 1996: 97. Type species: Haplanister rather straight; apex straight; lateral depressions absent; crypticus Moore, 1996, by monotypy. anterior angles feebly developed, angulate; posterior an- Description. Body length: 3.5–4.1 mm. Forebody (head gles strongly developed, rectangular; basal foveae deep, and thorax) without sparse setiferous micropores dorsally. wide; punctuation strongly developed (in basal foveae). Head. Mandibles short, strongly curved forward, blunt Apex of prosternal lobe with 5–6 long setae. Metepisterna apically. Labrum strongly transverse; apex straight medi- longer than wide. Legs. Male mesotarsi unmodified. ally. Eyes moderately large, convex, reaching buccal fissure Metatarsomere 1 as long as metatarsomeres 2+3. Elytra. ventrally. Tempora not inflated. Frons with clypeo-ocular 78 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) prolongations incomplete toward eyes. Antennal pubes- mum width. Thorax. Pronotum (Fig. 163) suborbicular, cence starting from basal third of antennomere 3. Mentum widest before middle; sides converging toward base, not with a tooth medially, moderately shorter than lateral lobes. sinuate; base strongly convex; apex rather straight; lateral Mentum and submentum separated by complete trans- depressions absent; anterior angles moderately developed, verse suture. Paraglossae longer than ligula. Palpi with last obtuse; posterior angles feebly developed, broadly rounded; segment fusiform, not truncate apically, with sparse, mod- basal foveae shallow, ill-defined; punctuation feebly devel- erately long pubescence; penultimate segment of labial palpi oped (in basal foveae). Metepisterna longer than wide. bisetose on anterior margin. Thorax. Pronotum Elytra. Widest about middle. Shoulders strongly devel- suborbicular; base convex, moderately narrower than elytral oped, rounded, without a tooth. Subapical sinuations ab- base; lateral beads complete; anterior and posterior beads sent. Sutural apices rounded. Scutellar striole present. In- incomplete medially. Scutellum visible. Apex of prosternal tervals impunctate, flat. Interval 3 with a setiferous punc- lobe glabrous. Legs. Metafemora with 2 long setae on pos- ture behind middle. Aedeagus (Fig. 80). Lateral view: as terior margin. Male protarsi and mesotarsi unmodified, for genus. Dorsal view: as for genus; dorsal membranous neither dilated laterally nor biseriately pubescent ventrally. area very wide. Segment 4 of protarsi and mesotarsi of both sexes without Material examined. 258 non-type specimens (AMNZ, membranous laminae. Tarsi glabrous dorsally; ITNZ, JNNZ, LUNZ, MONZ, NZAC, OMNZ, PHNZ, metatarsomere 5 pubescent (with 2 setae) ventrally; UCNZ). metatarsomere 1 shorter than metatarsomeres 2+3. Elytra. Geographic distribution (Map p. 152). North Island: Interneurs incomplete basally and laterally. Rows of AK, BP, CL, GB, HB, ND, RI, TK, TO, WA, WI, WN, setiferous punctures absent on intervals 3, 5, and 7, and on WO. South Island: MC, NN, SC. Offshore Islands: CH. interneur 2. Umbilicate setiferous series of interval 9 sepa- Adventive species of unknown origin. First New Zealand rated into 2 major groups, with posterior group continu- records: Napier, HB, 1965 (MONZ); Palmerston North, ous. Abdomen. Ventrites 2+3 of male without a setiferous WI, 1974 (Moore, 1996: 98). Well established. fovea. Ventrites 5+6 of both sexes without short setae, with paired ambulatory setae only. Aedeagus (Fig. 80). Ecology. Lowland, montane, subalpine, alpine. Forests Lateral view: strongly arcuate, especially stout and small (broadleaf, podocarp, beech). Gardens, golf courses, parks, (sabot-shaped). Dorsal view: symmetrical (with ostium pastures, fields, and alpine meadows. Shaded or open dorsal, not deflected laterally); dorsal membranous area ground. Usually nocturnal; sometimes active in the sun- wide, extending almost to basal bulb; apical disc absent. shine; generally sheltering during the day in leaf litter and Internal sac unarmed. in moss along logs and fallen trees, also under the loose bark of trees, in rotten wood, and among plant debris. Geographic distribution. New Zealand (apparently Gregarious. Biology. Seasonality: September–April, July. adventive). Tenerals: late January. Predators: Starlings (frequent en- References. Moore, 1996: 97-100 (taxonomy); Larochelle emies). Dispersal power. Elytra free along suture. & Larivière, 2001: 127 (catalogue). Macropterous. Frequent flier. Regular in seashore drift Remarks. A highly distinctive genus amongst the material, which indicates flight ability. Moderate runner. Stenolophina, with suborbicular pronotum, incomplete Good burrower. Regular climber (on plants, shrubs, and elytral interneurs, and sabot-shaped aedeagus. trees). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking leaf lit- ter, light trapping, turning seashore drift material, using Haplanister crypticus Moore, 1996 A malaise traps, pitfall trapping. Figures 80, 111, 163, 219; Map p. 152 References. Kuschel, 1990: 24, 40 (as Haplaner sp., dis- Haplanister crypticus Moore, 1996: 98. Type locality: Hast- tribution, ecology, biology, dispersal power); Townsend, ings, HB (although an adventive species). 1994: 9, 11 (as Haplaner sp., distribution, ecology); Moore Description. Body length: 3.5–4.1 mm. Slightly convex. 1996: 97–99 (distribution, ecology); Emberson, 1998: 30 Dark brown; base of antennae, maxillary palpi, femora, (distribution, ecology, biology); Larochelle & Larivière, and posterior 1/2 of tibiae light red. Generally glabrous and 2001: 127 (taxonomy, distribution, ecology, biology, dis- smooth. Microsculpture isodiametric. Shiny; pronotum persal power). and elytra with bronze lustre. Head. Moderately large, narrower across eyes than pronotal apex; flat anteriorly, slightly convex posteriorly. Antennae short, not reaching pronotal base; antennal scape about 2× longer than its maxi- Fauna of New Zealand 53 79

Genus Kiwiharpalus new genus E Kiwiharpalus townsendi new species E Type species. Kiwiharpalus townsendi new species, by present Figures 112, 164, 220; Map p. 152 designation. Kiwiharpalus townsendi Larochelle & Larivière, new spe- Description. Body length: 3.0–3.5 mm. Forebody (head cies. Holotype: female (NZAC) labelled “THREE KINGS and thorax) without sparse setiferous micropores dorsally. IS NZ, Princes I Hinemoa Nov 1983 C. F. Butcher (typed) Head. Mandibles very long (about 6× their maximum / Nest of Larus novaehollandiae scopulinus 83[forward width), slightly curved forward, acute apically. Labrum slash]131 (typed) / HOLOTYPE [male symbol] Kiwiharpalus townsendi Larochelle & Larivière, 2004 moderately transverse; apex straight medially. Eyes strongly (red label; typed).” Paratypes: 2 females (1 AMNZ, 1 reduced, flat, consisting of obliterated facets, narrowly sepa- × NZAC) from the same locality as the holotype, bearing rated from buccal fissure ventrally (by about 0.7 maxi- blue paratype labels. mum width of antennal scape). Tempora not inflated. Frons Description. Body length: 3.0–3.5 mm. Moderately con- with clypeo-ocular prolongations incomplete toward eyes. vex. Depigmented, testaceous; palpi partly greyish brown. Antennal pubescence starting from basal 1/3 of antennomere Generally glabrous and punctate (with sparse micropores) 3. Mentum with a tooth medially, moderately shorter than dorsally. Microsculpture absent. Shiny, without metallic lateral lobes. Mentum and submentum separated by com- lustre. Head. Big, although narrower across eyes than plete transverse suture. Paraglossae longer than ligula. Palpi pronotal apex; flat anteriorly, convex posteriorly. Anten- with last segment elongate-triangular, not truncate apically, nae rather long, reaching basal 1/3 of elytra; antennal scape with sparse, moderately long pubescence; penultimate seg- about 2× longer than its maximum width. Thorax. Pronotum ment of labial palpi bisetose on anterior margin. Thorax. (Fig. 164) quadrate (as wide as long), widest before middle; Pronotum quadrate (as wide as long); base rather straight, sides converging toward base, not sinuate; apex rather much narrower than elytral base; lateral beads complete; straight; lateral depressions widening posteriorly; anterior anterior and posterior beads incomplete medially. Scutellum angles feebly developed, obtuse; posterior angles moder- visible. Apex of prosternal lobe glabrous. Legs. Metafemora ately developed, rounded; basal foveae deep, wide; punc- with 2 long setae on posterior margin. Dilatation and ven- tuation strongly developed. Metepisterna longer than wide. tral vestiture of male pro- and mesotarsi unknown (only Elytra. Widest about middle. Shoulders strongly devel- females seen). Segment 4 of protarsi and mesotarsi with- oped, rounded, without a tooth. Subapical sinuations rather out membranous laminae. Tarsi glabrous dorsally (except strong. Sutural apices angulate. Scutellar striole absent. tarsomere 5); metatarsomere 5 pubescent (with 2 setae) Interneurs shallow, unevenly impressed, impunctate. In- ventrally; metatarsomere 1 as long as metatarsomeres 2+3. tervals sparsely punctate, flat. Interval 3 without setiferous Elytra. Interneurs complete. Rows of setiferous punc- puncture behind middle. Aedeagus. No male seen. tures absent on intervals 3, 5, and 7, and on interneur 2. Umbilicate setiferous series of interval 9 separated into 2 Material examined. 3 type specimens (AMNZ, NZAC). major groups, with posterior group further divided into 2 Geographic distribution (Map p. 152). Offshore Islands: subgroups. Abdomen. Condition of ventrites 2+3 of male TH–Princes Islands. unknown (only females seen). Ventrites 5+6 without short Ecology. Lowland. In a nest of red-billed gull (Larus setae, with paired ambulatory setae only. Aedeagus. No novaehollandiae scopulinus). Biology. Seasonality: No- male seen. vember. Dispersal power. Elytra fused along suture. Geographic distribution. New Zealand (endemic; Three Subapterous. Slow runner (inferred from body morphol- Kings Islands). ogy). Remarks. The generic name is derived from kiwi (the most Remarks. The strongly reduced eyes, depigmented and primitive of New Zealand birds and a major national sym- flattened body, and long pubescence suggest subterranean bol) and Harpalus (the type genus of the tribe Harpalini). behaviour similar to that of Anillina (Bembidiini) that live This monotypic genus appears to be a genetically highly deep in thick leaf litter and/or in soil fissures. This species distinctive taxon with its dorsal surface punctate through- is named after J. I. Townsend (Levin) for his contribution out and without microsculpture, its quadrate pronotum, to the building of important reference collections of New its small body length (3.0–3.5 mm), and its geographic Zealand carabids. isolation on the Three Kings Islands. 80 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Genus Pholeodytes Britton, 1962 E References. Britton, 1962: 665–668 and 1964a: 631 (tax- Pholeodytes Britton, 1962: 665. Type species: Pholeodytes onomy); Noonan, 1976: 27 (taxonomy); Larochelle & townsendi Britton, 1962, by monotypy. Larivière, 2001: 127–128 (catalogue). Description. Body length: 6.0–8.3 mm. Forebody (head Remarks. In addition to characters of the frons, labrum, and thorax) without sparse setiferous micropores dorsally; mentum, pronotum, and elytra, the presence of 2 genital much narrower than elytra (contrary to other Harpalini swellings on the aedeagus of Pholeodytes species sets this genera, except Syllectus). Antennae and legs very long (con- genus apart from all other Stenolophina genera. This genus trary to other Harpalini genera, except Syllectus). Head. only occurs in caves in the Northwest Nelson (NN) region. Mandibles very long (about 5× their maximum width), See also Remarks under Syllectus. slightly curved forward, acute apically. Labrum moder- ately transverse; apex convex medially. Eyes strongly re- duced, flat, consisting of obliterated facets, narrowly sepa- Key to species of Pholeodytes rated from buccal fissure ventrally (by about 0.8× maxi- mum width of antennal scape). Tempora not inflated. 1 Elytral interneurs coarsely punctate (Fig. 225). Pronotum Frons without clypeo-ocular prolongations. Antennal (Fig. 169): basal foveae very deep and wide ...... pubescence starting from basal 1/4 of antennomere 3...... (p. 83)... helmorei new species Mentum with a tooth medially, moderately longer than — Elytral interneurs finely punctate (Fig. 224). Pronotum lateral lobes. Mentum and submentum separated by com- (Fig. 168): basal foveae moderately deep, ill-defined plete transverse suture. Paraglossae longer than ligula...... 2 Palpi with last segment fusiform, not truncate apically, 2(1) Pronotum (Fig. 165): apex much wider than base; glabrous; penultimate segment of labial palpi bisetose on anterior angles subrectangular. Elytral interneurs deep × anterior margin. Thorax. Pronotum very long (almost 2 between base and apex (Fig. 221). Aedeagus (Fig. 81) longer than wide); base straight, much narrower than with genital swellings hook-like ...... elytral base; lateral beads complete; anterior and poste- ...... (p. 81)... palmai new species rior beads absent (as in Syllectus gouleti). Scutellum vis- — Pronotum (Fig. 168): apex only slightly wider than ible. Apex of prosternal lobe glabrous. Legs. Metafemora base; anterior angles rounded. Elytral interneurs shallow with 3–4 long setae on posterior margin. Male protarsi (Fig. 224). Aedeagus with genital swellings not hook- dilated laterally and biseriately pubescent ventrally. Male like ...... 3 mesotarsi slightly dilated laterally, not biseriately pubes- cent ventrally. Segment 4 of protarsi and mesotarsi of 3(1) Pronotum (Fig. 166): posterior angles acute; sides both sexes with 2 membranous laminae (projecting later- strongly rounded in apical 1/2. Elytra subelliptical ally and anteriorly to about 2/3 the length of apical seg- (narrower at base; Fig. 222). Aedeagus (Fig. 82) with ment; as in Syllectus). Tarsi pubescent (with few setae) genital swellings subtriangular ...... dorsally; metatarsomere 5 pubescent (with 7-8 setae) ...... (p. 81)... cerberus Britton ventrally; metatarsomere 1 as long as metatarsomeres 2+3. — Pronotum (Fig. 167–168): posterior angles rounded; Elytra. Interneurs incomplete, consisting of rows of punc- sides moderately rounded in apical 1/2. Elytra elliptical tures (contrary to other Stenolophine genera). Rows of (base as wide as apex; Fig. 223–224 ). Aedeagus with setiferous punctures absent on intervals 3, 5, and 7, and genital swellings not subtriangular ...... 4 on interneur 2. Umbilicate setiferous series of interval 9 4(3) Pronotum (Fig. 168): posterior angles obtusely divided into two major groups, with posterior group con- rounded; sides barely sinuate in basal 1/2. Aedeagus tinuous. Abdomen. Ventrites 2+3 of male without a (Fig. 84): genital swellings subelliptical ...... setiferous fovea. Ventrites 5+6 of both sexes without short ...... (p. 82)... townsendi Britton setae, with paired ambulatory setae only. Aedeagus. Lat- eral view: strongly arcuate. Dorsal view (Fig. 81–84): — Pronotum (Fig. 167): posterior angles acutely rounded; symmetrical (with ostium dorsal, not deflected laterally); sides moderately sinuate in basal 1/2. Aedeagus (Fig. dorsal membranous area wide (with 2 genital swellings), 83): genital swellings subrectangular ...... not extending to basal bulb; apical disc absent. Internal ...... (p. 82)... nunni new species sac unarmed. Geographic distribution. New Zealand (endemic; South Island). Fauna of New Zealand 53 81

Pholeodytes palmai new species E Pholeodytes cerberus Britton, 1964 E Figures 81, 165, 221; Map p. 153 Figures 82, 166, 222; Map p. 152 Pholeodytes palmai Larochelle & Larivière, new species. Pholeodytes cerberus Britton, 1964a: 631. Holotype: male Holotype: male (NZAC) labelled “Pluto’s Retreat Cave (NZAC) labelled “Type (circular red-bordered label; typed) Kaihoka N.W. Nelson 16.1.75 S.-I. Uéno (hand-written) / Fenian Crk. Cave Oparara, Karamea 27.4.63 R Main. / HOLOTYPE [male symbol] Pholeodytes palmai (hand-written) / HOLOTYPE [male symbol] Pholeodytes Larochelle & Larivière, 2004 (red label; typed).” cerberus mihi (hand-written) E.B. Britton det. 1963 Paratype: 1 male (MONZ) from the same locality as the (typed, except for number 3).” Perfect condition. holotype, bearing blue paratype label. There are also 2 paratypes in NZAC. Description. Body length: 6.5–7.0 mm. Slightly convex. Description. Body length: 6.0–7.6 mm. Slightly convex. Depigmented (appearing pale in colour), pale yellowish Depigmented (appearing pale in colour), reddish; anten- (teneral condition). Generally glabrous and smooth. nae, palpi, and legs yellowish brown. Generally glabrous Microsculpture weak, isodiametric on head, very trans- and smooth. Microsculpture isodiametric and rather strong verse (with microlines) on pronotum and elytra. Shiny, on head, very transverse (with microlines) and vestigial on without metallic lustre. Head. Narrow, although as wide pronotum and elytra. Shiny, without metallic lustre. Head. across eyes as pronotal apex; flat anteriorly, slightly con- Narrow, although as wide across eyes as pronotal apex; vex posteriorly. Antennae very long, reaching middle of flat anteriorly, slightly convex posteriorly. Antennae very elytra; antennal scape elongate, about 3× longer than its long, reaching middle of elytra; antennal scape elongate, maximum width. Thorax. Pronotum (Fig. 165) widest be- about 3× longer than its maximum width. Thorax. Pronotum fore middle; sides feebly rounded in apical 1/2, slightly (Fig. 166) widest before middle; sides strongly rounded in sinuate in basal 1/2; base straight, much narrower than apical 1/2, strongly sinuate in basal 1/2; base straight, much elytral base; apex straight, much wider than base (contrary narrower than elytral base; apex straight; lateral depres- to other Pholeodytes species); lateral depressions widen- sions widening posteriorly; anterior angles feebly devel- ing posteriorly; anterior angles moderately developed, oped, rounded; posterior angles strongly developed, acute, subrectangular; posterior angles strongly developed, projecting laterally; basal foveae moderately deep, ill-de- subrectangular, not projecting laterally; basal foveae mod- fined; punctuation feebly developed. Metepisterna longer erately deep, ill-defined; punctuation feebly developed. than wide. Elytra. Subelliptical (narrower at base). Widest Metepisterna longer than wide. Elytra. Elliptical (base as behind middle. Shoulders feebly developed, without a tooth. wide as apex). Widest about middle. Shoulders feebly de- Subapical sinuations feeble. Sutural apices angulate. veloped, without a tooth. Subapical sinuations feeble. Scutellar striole absent. Interneurs shallow, finely punc- Sutural apices angulate. Scutellar striole absent. Interneurs tate. Intervals impunctate, flat. Interval 3 without setiferous deep between base and apex, finely punctate. Intervals puncture behind middle. Aedeagus (Fig. 82). Dorsal view: impunctate, slightly convex. Interval 3 without setiferous as for genus; subapical area long; apex moderately wide, puncture behind middle. Aedeagus (Fig. 81). Dorsal view: rounded; genital swellings subtriangular. as for genus; subapical area short; apex moderately wide, Material examined. 9 specimens, including types (ITNZ, rounded; genital swellings hook-like. LUNZ, NZAC). Material examined. 3 specimens, including types (ITNZ, Geographic distribution (Map p. 152). South Island: MONZ, NZAC). NN–Fenian Creek Cave, Oparara. Honeycomb Hill Caves Geographic distribution (Map p. 153). South Island: (east of Karamea Bluff). Ida Cave, Oparara. Oparara. Star- NN–Pluto’s Retreat Cave, Kaikoka, West Haven Inlet. Draft Cave, Oparara. Ecology. Lowland. A cave (troglobitic species). Biology. Ecology. Lowland. Caves (troglobitic species). Found in Seasonality: unknown (adults). Tenerals: October, Janu- low roof crevice and on stalactite ceiling; on dry gypsum ary. Dispersal power. Elytra fused along suture. sand at some distance from dripping or wet areas. Biology. Subapterous. Fast runner. Collecting techniques. Using Seasonality: March–April. Tenerals: September, January, a head lamp or torch. March–April. Occasionally infested with fungi Remarks. This species is named after our friend and col- (Laboulbeniales). Dispersal power. Elytra fused along league Ricardo L. Palma (Museum of New Zealand Te suture. Subapterous. Fast runner. Good climber. Collect- Papa Tongarewa, Wellington) for his special help and en- ing techniques. Using a head lamp or torch. couragement in our entomological studies. References. Britton, 1964a: 633 (distribution, ecology); May, 1972: 575 (ecology); Johns, 1991: 20 (distribution); Townsend, 1997: 17 (distribution, ecology); Larochelle & 82 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Larivière, 2001: 127-128 (taxonomy, distribution, ecology, except December. Tenerals: January, March, June. Dis- biology, dispersal power). persal power. Elytra fused along suture. Subapterous. Fast runner. Good climber. Collecting techniques. Using a head lamp or torch. Pholeodytes nunni new species E Remarks. This new taxon corresponds to Britton’s (1964a) “Pholeodytes cerberus var. A” from Takaka Hill, illustrated Figures 83, 167, 223; Map p. 152 Pholeodytes nunni Larochelle & Larivière, new species. and characterised, although not officially described as a Holotype: male (NZAC) labelled “Council Cave Motupipi subspecies. This species is named after John Nunn Takaka 14.6.73 L McRae (hand-written) / HOLOTYPE (Dunedin) for his contribution to the building of important [male symbol] Pholeodytes nunni Larochelle & Larivière, reference collections of New Zealand carabids. 2004 (red label; typed).” Paratypes: 7 males (1 CMNZ, 2 LUNZ, 3 NZAC, 1 OMNZ) and 6 females (1 CMNZ, 1 LUNZ, 3 NZAC, 1 OMNZ) from the same locality as the Pholeodytes townsendi Britton, 1962 E holotype, bearing blue paratype labels. Figures 84, 113, 168, 224; Map p. 153 Description. Body length: 7.0–8.0 mm. Slightly convex. Pholeodytes townsendi Britton, 1962: 666. Holotype: male Depigmented (appearing pale in colour), reddish brown; (NZAC) labelled “Type (circular red-bordered label; typed) antennae, palpi, and legs pale yellow. Generally glabrous / Twin Forks Cave Paturau NELSON 30.1.61. (hand- and smooth. Microsculpture isodiametric and rather strong written) J.I. Townsend . (typed) / J.I. Townsend Collec- on head, very transverse (with microlines) and weak on tion (hand-written, in red ink) / HOLOTYPE [male sym- bol] Pholeodytes townsendi mihi (hand-written) E.B. pronotum and elytra. Shiny, without metallic lustre. Head. Britton. det. 1961 (typed, except for number 1 in 1961).” Narrow, although as wide across eyes as pronotal apex; Perfect condition. flat anteriorly, slightly convex posteriorly. Antennae very There are also 32 paratypes in NZAC (29) and BMNH (3). long, reaching middle of elytra; antennal scape elongate, Description. Body length: 7.0–8.0 mm. Slightly convex. × about 3 longer than its maximum width. Thorax. Pronotum Depigmented (appearing pale in colour), yellowish brown. (Fig. 167) widest before middle; sides moderately rounded Generally glabrous and smooth. Microsculpture in apical 1/2, moderately sinuate in basal 1/2; base straight, isodiametric and rather strong on head, very transverse narrower than elytral base; apex straight; lateral depres- (with microlines) and weak on pronotum and elytra. Shiny, sions widening posteriorly; anterior angles feebly devel- without metallic lustre. Head. Narrow, although as wide oped, rounded; posterior angles strongly developed, acutely across eyes as pronotal apex; flat anteriorly, slightly con- rounded, projecting laterally; basal foveae moderately deep, vex posteriorly. Antennae very long, reaching middle of ill-defined; punctuation feebly developed. Metepisterna elytra; antennal scape elongate, about 3× longer than its longer than wide. Elytra. Elliptical (base as wide as apex); maximum width.Thorax. Pronotum (Fig. 168) widest be- elytra broader than in cerberus. Widest about middle. Shoul- fore middle; sides moderately rounded in apical 1/2, barely ders feebly developed, without a tooth. Subapical sinuations sinuate in basal 1/2; base straight, much narrower than feeble. Sutural apices angulate. Scutellar striole absent. elytral base; apex straight; lateral depressions widening Interneurs shallow, finely punctate. Intervals impunctate, posteriorly; anterior angles feebly developed, rounded; flat. Interval 3 without setiferous puncture behind middle. posterior angles strongly developed, obtusely rounded, Aedeagus (Fig. 83). Dorsal view: as for genus; subapical not projecting laterally; basal foveae moderately deep, ill- area long; apex moderately wide, rounded; genital swell- defined; punctuation feebly developed. Metepisterna longer ings subrectangular. than wide. Elytra. Elliptical (base as wide as apex). Widest Material examined. 83 specimens, including types (ITNZ, about middle. Shoulders feebly developed, without a tooth. LUNZ, NZAC). Subapical sinuations feeble. Sutural apices angulate. Geographic distribution (Map p. 152). South Island: Scutellar striole absent. Interneurs shallow, finely punc- NN–Council Cave, Motupipi, Takaka. Ervin’s Cave, tate. Intervals impunctate, flat. Interval 3 without setiferous Takaka. Gorge Creek Cave, Takaka Valley. Green Link Cave, puncture behind middle. Aedeagus (Fig. 84). Dorsal view: Canaan Road, Takaka Hill. Kerry-John Cave, Barron Flat, as for genus; subapical area long; apex very wide, truncate; Takaka. Sims Cave, Takaka Valley. Water Supply Cave, genital swellings subelliptical. Pohara. Weka Cave, Upper Takaka. Worm’s Surprise Cave, Material examined. 89 specimens, including NZAC types Barron Flat, Takaka. (BMNH, ITNZ, JNNZ, LUNZ, NZAC, UCNZ). Ecology. Lowland, montane. Caves (troglobitic species): Geographic distribution (Map p. 153). South Island: on calcite walls. Biology. Seasonality: throughout the year, NN-17 caves between Paturau and Heaphy Track areas. Fauna of New Zealand 53 83

Ecology. Lowland. Caves (troglobitic species): in dry gyp- Geographic distribution (Map p. 152). South Island: sum sand at some distance from dripping or wet areas. NN–Coal Flat Cave [=Eggers Cave, J. I. Townsend, per- Biology. Seasonality: September–June. Tenerals: Septem- sonal communication.] ber, November, March–April. Predacious. Food: a small Ecology. Lowland. Cave (troglobitic species). Biology. weta. Dispersal power. Elytra fused along suture. Seasonality: November, July. Subapterous. Fast runner. Good climber. Collecting tech- Dispersal power. Elytra fused along suture. Subapterous. niques. Using a head lamp or torch. Fast runner. Collecting techniques. Using a head lamp or References. Britton, 1962: 666, 668 (distribution, biol- torch. ogy); Townsend, 1963: 96 (distribution, ecology); May, Remarks. This species is named after our friend and col- 1972: 575 (ecology); Johns, 1991: 20 (distribution); league Desmond W. Helmore (Landcare Research, Auck- Townsend, 1997: 17 (distribution, ecology); Larochelle & land) for supporting our entomological studies and for his Larivière, 2001: 128 (taxonomy, distribution, ecology, bi- exceptional talent and contribution as illustrator of New ology, dispersal power). Zealand insects.

Pholeodytes helmorei new species E Figures 169, 225; Map p. 152 Pholeodytes helmorei Larochelle & Larivière, new species. Holotype: female (NZAC) labelled “Coal Flat Cave W. BIBLIOGRAPHY of New Creek Buller, 1.11.75 J. I. Townsend (hand-writ- ten) / HOLOTYPE [female symbol] Pholeodytes Acorn, J. H.; Ball, G. E. 1991: The mandibles of some helmorei Larochelle & Larivière, 2004 (red label; typed).” adult ground beetles: structure, function, and the Paratype: 1 female (NZAC) from the same locality as evolution of herbivory (Coleoptera: Carabidae). the holotype, bearing blue paratype labels. Canadian Journal of Zoology 69: 638–650. Description. Body length: 8.0–8.3 mm. Slightly convex. Andrewes, H. E. 1924: On the Oriental Carabidae of the Depigmented (appearing pale in colour), reddish; anten- “Reise Novara”. Transactions of the Royal nae, palpi and legs yellowish brown. Generally glabrous Entomological Society of London 1923: 459–468. and smooth. Microsculpture isodiametric and rather strong ——— 1935: On the genotypes of British Carabidae. — on head, very transverse (with microlines) and weak on II. Annals and Magazine of Natural History (10) 16: pronotum and elytra. Shiny, without metallic lustre. Head. 12–25. Narrow, although as wide across eyes as pronotal apex; Arndt, E. 1998: Phylogenetic investigation of Carabidae flat anteriorly, slightly convex posteriorly. Antennae very (Coleoptera) using larval characters. Pp. 171–190. In: long, reaching middle of elytra; antennal scape elongate, Ball, G. E.; Casale, A.; Taglianti, A. V. (Editors). 1998: about 3× longer than its maximum width. Thorax. Pronotum Phylogeny and classification of Caraboidea (Fig. 169) widest before middle; sides moderately rounded (Coleoptera: Adephaga). Proceedings of a Symposium in apical 1/2, not sinuate, obliquely converging in basal 1/2; (28 August, 1996, Florence, Italy). XX International base straight, much narrower than elytral base; apex straight; Congress of Entomology. Atti Museo Regionale di lateral depressions widening posteriorly; anterior angles Scienze Naturali, Torino. 543 pp. feebly developed, rounded; posterior angles strongly de- veloped, subrectangular, not projecting laterally; basal Ball, G. E.; Bousquet, Y. 2001: 6. Carabidae Latreille, foveae very deep, wide; punctuation feebly developed. 1810. Pp. 32–132. In: Arnett, R. H. Jr.; Thomas, M. Metepisterna longer than wide. Elytra. Subelliptical (nar- C. (Editors). American beetles. Vol. 1. CRC Press, rower at base). Widest behind middle. Shoulders feebly Boca Raton, London, New York, Washington D.C. developed, without a tooth. Subapical sinuations feeble. 443 pp. Sutural apices angulate. Scutellar striole absent. Interneurs Barratt, B. I. P.; Patrick, B. H. 1987: Insects of the snow moderately deep, coarsely punctate; interneur 8 deeply tussock grassland on the East Otago Plateau. New impressed (more so than in other Pholeodytes species). Zealand Entomologist 10: 69–98. Intervals impunctate, slightly convex. Interval 3 without Basilewsky, P. 1950: Révision générale des Harpalinae setiferous puncture behind middle. Aedeagus. No male d’Afrique et de Madagascar (Coleoptera Carabidae). seen. Première Partie. Annales du Musée du Congo Belge Material examined. 3 specimens, including types (ITNZ, Tervuren (Belgique). Séries 8º, Sciences Zoologiques NZAC). 6: 1–283. 84 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

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——— 1878: Les harpaliens de l’Australie d’après la synonymis, locis, descriptionibus, observationibus. collection de M. le Comte de Castelnau et la mienne. Kortii, Flensburgi et Lipsiae. XXXII + 832 pp. Annali del Museo Civico di Storia Naturale di Genova ——— 1787: Mantissa insectorum, sistens eorum species 12: 475–517. nuper detectas adiectis characteribus genericis, Crosby, T. K.; Dugdale, J. S.; Watt, J. C. 1976: Recording differentiis specificis, emedationibus, observationibus. specimen localities in New Zealand: an arbitrary Tom. I. Proft, Hafniae. XX + 348 pp. system of areas and codes defined. New Zealand ——— 1798: Supplementum entomologiae systematicae. Journal of Zoology 3: 69 (with separate map). Proft et Storch, Hafniae. II + 572 pp. ——— 1998: Area codes for recording specimen localities Fauvel, A. 1882: Les coléoptères de la Nouvelle-Calédonie in the New Zealand subregion. New Zealand Journal et dépendances avec descriptions, notes et synonymies of Zoology 25: 175–183. nouvelles. Revue d’Entomologie 1: 217–236, 241– Csiki, E. 1932: Carabidae: Harpalinae VI. Pars 121: 1023– 261, 265–274. 1278. In: Junk, W.; Schenkling, S. (Editors), Ganglbauer, L. 1892: Die Käfer von Mitteleuropa. Erster Coleopterorum Catalogus. Junk, Berlin. Band. Familienreihe Caraboidea. Gerold, Wien. III + Darlington, P. J. Jr. 1956: Australian carabid beetles. III. 557 pp. Notes on the Agonini. Psyche 63: 1–10. Gemminger, M.; Harold, E. von. 1868: Catalogus ——— 1968: The carabid beetles of New Guinea part III. coleopterorum hucusque descriptorum synonymicus Harpalinae (continued): Perigonini to Pseudomorphini. et systematicus. Tomus I. Cicindelidae-Carabidae. Bulletin of the Museum of Comparative Zoology 137: Gummi, Monachii. XXXVI + 424 pp + 8 unnumbered 1–253. page. Index. ——— 1971: Modern taxonomy, reality, and usefulness. Germar, E. F. 1848: Beiträge zur Insectenfauna von Systematic Zoology 20(3): 341–365. Adelaide. Linnaea Entomologici 3: 153–247. DeGeer, C. 1774: Mémoires pour servir à l’histoire des Goulet, H. 1974: Classification of the North and Middle insectes. Tome quatrième. Hesselberg, Stockholm. XII American species of the genus Pelmatellus Bates + 456 pp. + 19 plates. (Coleoptera: Carabidae: Harpalini). Quaestiones Dejean, P. F. M. A. 1829: Species général des Coléoptères, Entomologicae 10: 80–102. de la collection de M. le Comte Dejean. Tome Guérin-Méneville, F. É. 1830: Insectes. Atlas. Planches I- quatrième. Méquignon-Marvis, Paris. VII + 520 pp. XXI. In: Duperrey, L. J., 1838, Voyage autour du Duftschmid, C. E. 1812: Fauna Austriae. Oder Monde, exécuté par ordre du roi, sur la Corvette de sa Beschreibung der österreichischen Insekten für Majesté, la Coquille, pendant les années 1822, 1823, angehende Freunde der Entomologie. Zweiter Theil. 1824, et 1825... Histoire naturelle: zoologie. Bertrand, Akademie Buchhandlung, Linz und Leipzig. VIII + Paris. 1830–1832 [Descriptions to the plates 311 pp. appeared in Guérin-Méneville, F. É., 1838, Crustacés, Emberson, R. M. 1998: The beetle (Coleoptera) fauna of arachnides et insectes, pages 57–302 in Duperrey, L. the Chatham Islands. New Zealand entomologist 21: J., 1838, Voyage autour du Monde, exécuté par ordre 25–64. du Roi, sur la Corvette de sa Majesté, la Coquille, ——— 2004: A European ground beetle, Harpalus tardus pendant les années 1822, 1823, 1824, et 1825... (Panzer) (Coleoptera: Carabidae) in Canterbury, and Zoologie, Volume 2, Partie 2, Division 1, Bertrand, the predisposition of European carabids to establish in Paris.] exotic temperate locations. New Zealand Entomologist Habu, A. 1973: Carabidae: Harpalini (Insecta: Coleoptera). 27: 39–42. Fauna Japonica. Keigaku Publishing Co., Tokyo. 430 Emden, F. I. van. 1953: The Harpalini genus Anisotarsus pp. Dejean (Col. Carab.). Annals and Magazine of Natural Harris, A. C. 1970: Coastal beetles of the Wanganui- History, Series 12 (6): 513–547. Manawatu area. Part 1. Environmental factors and Erichson, W. F. 1842: Beitrag zur Insecten-Fauna von zonation. Bulletin of Natural Sciences (Wellington) 1: Vandiemensland, mit besonderer Berüsichtigung der 45–58. geographischen Verbreitung der Insecten. Archiv für Hudson, G. V. 1934: New Zealand beetles and their larvae: Naturgeschichte 8: 83–287. an elementary introduction to the study of our native Fabricius, J. C. 1775: Systema entomologiae, sistens Coleoptera. Ferguson and Osborn, Wellington. 236 pp insectorum classes, ordines, genera, species, adiectis + 18 plates. 86 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

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Appendix A. Glossary of technical terms. facet — lens-like division of each compound eye. family — a category in the taxonomic hierarchy, adventive — not native; an organism carried into a that includes one or more genera or tribes of new habitat by natural means, or by man. common phylogenetic origin, separated from aeneous — with a copper or brass appearance. other such groups by a decided gap. allopatric — of or pertaining to taxa occupying dif- fovea — a small pit or depression. ferent and disjunct geographical areas. fusiform — spindle-shaped. genus — a category in the taxonomic hierarchy, alpine — of or pertaining to land located above the that includes one or more phylogenetically re- subalpine zone, characterised by grasslands, lated, and morphologically similar species. herb fields and screes, and reaching up to the geographic distribution — distribution related to summer snow line. geography, i.e., districts, regions. ambulatory setae — specialised pairs of setae occurring ventrally on the abdomen. glabrous — without hairs or setae (not pubescent). antennomere — each antennal segment. granivorous — eating grains or seeds. apex — end or extremity of a structure or organ. granulate (of microsculpture) — appearing covered apical — related to the apex. with small grains. appendages — antennae, palpi, and legs together. gregarious — living in groups or colonies. armed — displaying scales, spines, or teeth. hirsute — densely covered with long shaggy se- attenuate — gradually tapering toward the apex. tae. bead — a raised border. biostatus — the status of an organism based on holotype or type — the single specimen desig- its geographic origin relative to its occurrence nated or indicated as the type specimen of a in a particular region, e.g., endemic, native, species by the original author at the time of publication or the only specimen from which adventive. the original description was made. biseriately — disposed in two rows. hygrophilous — living in moist or wet environments. bisetose — with two setae. brachypterous — with abbreviated membranous indigenous — see native. wings, shorter than those of macropterous interneur —a longitudinal stria (impressed line) species but not vestigial like those of or row of punctures on the elytron. subapterous species; incapable of flight. interval — the space between two interneurs on buccal fissure — lateral mouth opening beneath the elytron. eye area. iridescent — displaying a rainbow-effect colora- clypeo-ocular prolongations — deep lines be- tion. tween the clypeus and the eyes. isodiametric (of microsculpture) — appearing cov- coastal — of or pertaining to the strip of land within ered with polygons of equal diameter. the influence of the sea. lamina — a thin flat scale-like structure. conchoid — shell-shaped. lectotype — type specimen selected from the cordate — heart-shaped. syntypes by a subsequent author in the ab- deflected — turned aside. sence of a holotype. depigmented — with weak pigmentation lowland — of or pertaining to land located below (appearing pale in colour). the montane zone and generally reaching up disc — dorsal central area of a body part. to the limit of rimu (Dacrydium cupressinum), dispersal power — capacity of self-dispersal. e.g., about 500 m in central New Zealand. distal — situated farthest from the centre. macropterous — with long or fully developed mem- branous wings. diurnal — active during the day. medially — situated in the middle. emarginate — having a notched edge. monophyletic — referring to a group of taxa con- endemic — restricted to a geographic area. taining all descendants from a single hypo- explanate — spread and flattened. thetical ancestral taxon. extralimital range — distribution of an organism monotypy — the situation when a nominal genus outside the limits of a specific geographic area or subgenus is established on the basis of a (e.g., outside New Zealand). single species (the type species by monotypy). 90 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) montane — of or pertaining to land located above stria — elytral interneur in the form of an impressed the lowland zone and reaching up to the tree longitudinal line. line. sub- (as a prefix) — rather, almost. native — occurring naturally in the area under con- subalpine — of or pertaining to land located above sideration. the tree line and characterised by a mountain nocturnal — active during the night. shrubland (e.g., of Olearia, Brachyglottis, and omnivorous — feeding on both animal and veg- Dracophyllum). etable matter. subapical sinuation — sinuation of the lateral bor- der of each elytron, near its apex. orbicular — circular or spherical. subapterous — with vestigial membranous wings original designation — the situation when the type (reduced to small wing buds). of a taxon (genus or subgenus) is designated supraorbital — above the eye. at the same time as the taxon is established (the type species by original designation). sutural — related to the suture. ostium — membranous opening of the aedeagus. sutural apex, plural apices (of elytron) — inner ovate — egg-shaped. apex of each elytron. palpomere — each segment of a palp (palpus). sutural interval (of elytron) — the first interval next pedunculate — stalked. to the suture. penultimate — next to the last (e.g., penultimate suture (of elytron) — line of contact between the segment, the segment next to the last one). elytra. phytophagous — feeding on plant material. synonym — one of two or more scientific names piceous — pitchy black or black with reddish tinge. applied to a single taxon. plurisetose — with 4 or more setae. syntype — any of two or more specimens on which predacious — eating live animals. the original description of a taxon was based when a holotype was not designated. pubescence (adj. pubescent) — covering of hair tarsomere — each segment of a tarsus. or setae. taxon, plural taxa — a taxonomic grouping of any punctate — marked with points or punctures. rank (e.g., a family, a genus, a species) includ- quadrate — square or nearly so. ing all its subordinate groups. rufous — reddish. teneral — a new or young adult, recently emerged, sexually immature, with softer or paler exoskel- scree — accumulation of loose stones on a slope. eton. scrobe (of mandible) — a lateral depression in the testaceous — reddish brown. wall of the mandible. transverse (of microsculpture) — appearing cov- scrubland — vegetation with dense cover and about ered with flattened or sublinear shapes. 1–2 metres tall. trisetose — with three setae. scutellar striole — short stria on each side of the troglobitic — living exclusively in caves. scutellum. troglophilous — living usually, but not exclusively seasonality — period(s) of the year when an ani- in caves. mal is active. type or name-bearing type — the specimen(s), setiferous — bearing seta(e). species or genus that serves as the objective shrubland — vegetation with sparse or moderate standard of reference determining the appli- cover and often taller than 2 metres. cation of a name to a taxon. spatulate — spoon-shaped. type locality — the precise geographical site where the type of a species or subspecies was col- species — a taxon of the rank of species, the cat- lected. egory below the genus in the taxonomic hier- type species — the species designated as the archy; naturally occurring populations with a type of a genus or subgenus. common heredity; groups of actually or poten- type specimen — a specimen (e.g., holotype, tially interbreeding populations which are lectotype, neotype) or one of a series of speci- reproductively isolated from other such groups. mens (syntypes) designated as the type of a spongily — in a sponge-like formation. species or subspecies. Fauna of New Zealand 53 91 umbilicate series— row of setiferous punctures vestigial — strongly reduced, almost obsolete or along interval 9 on the elytron. absent. uniperforated — appearing pierced with one hole. ventrite — each ventral segment of the abdomen. valid name — the name for a particular taxon that is correct according to the provisions of the Code of Zoological Nomenclature.

Appendix B. Geographical coordinates of main Hastings, HB ...... 3939/17651 localities. Coordinates should read as 00º00’S/ Heaphy Track, NN ...... 4052/17226 000º00’E . The two-letter area codes follow Crosby Hikurangi, ND ...... 3535/17417 et al. (1976, 1998). Honeycomb Hill, east of Karamea Bluff, NN ...... 4107/17211 Ahipara, ND ...... 3510/17309 Howick, AK ...... 3654/17456 Auckland, AK ...... 3651/17446 Inangahua, BR ...... 4152/17157 Banks Peninsula, MC ...... 4340/17245 Barron Flat, Takaka, NN ...... 4105/17246 Kaihoka [Lakes], NN ...... 4033/17236 Belmont, WN ...... 4111/17455 Kaimai Summit [=Saddle], BP ...... 3752/17555 Boulder Lake, NN ...... 4054/17235 Kaitoke [Regional Park], WN ...... 4105/17511 Brown Cow, Boulder Lake Track, NN 4045/17234 Kaituna Valley, MC ...... 4345/17241 Buller River, near Inangahua, BR ...... 4150/17153 Kapowairua, Spirits Bay, ND ...... 3426/17252 Bullock Creek, Paparoa National Park, BR ...... Kara, ND ...... 3543/17412 ...... 4206/17124 Karamea, NN ...... 4115/17206 Karamea Bluff, NN ...... 4131/17201 Calphurnia Peak, NN ...... 4054/17234 Kohuronaki, ND ...... 3429/17250 Canaan [Road], Takaka, NN ...... 4057/17254 Kowhai River, KA ...... 4224/17338 Canterbury, NC/MC ...... 4330/17200 Capleston, BR ...... 4204/17155 Lake Coleridge, MC ...... 4322/17132 Charleston, BR...... 4201/17133 Lake Ohia, ND ...... 3458/17322 Christchurch, MC...... 4332/17238 Lincoln, MC...... 4338/17229 Clarence River mouth, KA ...... 4209/17355 Coal Flat, west of New Creek, NN..... 4148/17156 Maitai Valley, NN ...... 4116/17317 Coroglen, CL ...... 3655/17541 Maketu [Stream], Hunua Ranges, AK 3708/17500 Coromandel Range, CL ...... 3705/17542 Manawatu Gorge, RI/WN ...... 4018/17546 Mangakahia Valley, ND ...... 3537/17350 Donnelly’s Crossing, ND ...... 3542/17336 Mangamuka Gorge, ND ...... 3519/17332 Dunedin, DN ...... 4552/17030 Matakana, AK ...... 3621/17443 Maumaupaki, CL ...... 3658/17534 Epsom, AK ...... 3654/17446 McClean’s Island, MC ...... 4328/17228 Fantail Creek, Moehau Range, CL .... 3631/17520 Morere Springs Scenic Reserve, GB . 3857/17747 Forty Mile Bush [Masterton–Woodville], WA ...... Motueka, NN ...... 4106/17300 ...... 4035/17541 Motupipi, Takaka, NN ...... 4051/17251 Fox River, BR ...... 4203/17130 Mount Arthur, NN ...... 4116/17241 Mount Bossu, Banks Peninsula, MC 4350/17254 Glasgow Range, NN...... 4136/17204 Mount Domett, NN ...... 4104/17219 Gorge Creek, Takaka Valley, NN...... 4056/17251 Mount Hutt, MC ...... 4328/17132 Gowanbridge, Buller Gorge, NN ...... 4143/17233 Mount Messenger, TK ...... 3854/17456 Great Island, TH ...... 3410/17208 Mount Pleasant, Christchurch, MC .... 4120/17358 92 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Napier, HB ...... 3930/17654 Sandy Bay, Te Paki Coastal Park, ND 3426/17242 Nelson, NN...... 4113/17317 South East Island, CH ...... 4420/17610W New Creek, NN ...... 4147/17201 South West Island, TH ...... 3411/17204 Nile River, BR ...... 4156/17128 Spirits Bay, ND ...... 3427/17247 North Cape, ND ...... 3425/17303 Spreydon, MC ...... 4333/17237 North East Island, CH ...... 3408/17210W Swanson, AK ...... 3652/17434

Okaihau, ND ...... 3519/17346 Taieri, DN ...... 4605/17011 Okauia, WO ...... 3747/17550 Takaka, NN ...... 4051/17248 Omahuta Forest, ND...... 3514/17337 Takaka Hill, NN ...... 4053/17249 Oparara, NN ...... 4113/17209 Takaka Valley, NN ...... 4106/17243 Orongorongo Valley, WN ...... 4125/17454 Tangihua Range, ND ...... 3553/17407 Otaki Forks [=Gorge], WN ...... 4050/17515 Tapotupotu Bay, ND ...... 3426/17242 Tapu, CL ...... 3659/17530 Palmerston North, WI ...... 4022/17537 Te Koau, BP ...... 3735/17819 Pandora, ND ...... 3427/17247 Te Paki, ND ...... 3433/17247 Paparoa National Park [=Paparoa Range], BR .... The Sisters, CH ...... 4638/16915W ...... 4205/17133 Tiropahi Valley, BR ...... 4157/17126 Papatea, BP ...... 3740/17751 Titirangi Bay, SD ...... 4101/17409 Parakao, ND ...... 3543/17357 Tom Bowling Bay, ND ...... 3426/17257 Parua [Bay], ND ...... 3546/17427 Trounson Kauri Park, ND ...... 3544/17339 Paturau [Trig], NN ...... 4042/17229 Peraki [Saddle], Banks Peninsula, MC ...... Unuwhao, Spirits Bay, ND ...... 3426/17253 ...... 4349/17251 Upper Takaka, NN ...... 4102/17249 Pitt Island, CH ...... 4421/17609W Pohara, NN...... 4050/17253 Waiau River, Upper NC ...... 4243/17243 Port Ligar, SD ...... 4056/17359 Waimana Valley, BP ...... 3806/17702 Port Waikato, WO ...... 3724/17444 Waioeka Gorge, BP ...... 3810/17718 Princes Islands, TH ...... 3410/17202 Waipatiki Beach, HB ...... 3917/17658 Puketi Forest, ND ...... 3514/17346 Waipoua Forest, ND ...... 3539/17333 Punakaiki, BR ...... 4207/17120 Waipuna Stream, Spirits Bay, ND ...... 3426/17252 West Haven Inlet, NN ...... 4036/17235 Rangiputa, ND ...... 3452/17318 West Island, TH ...... 3411/17202 Ratapihipihi, TK ...... 3906/17403 Whanarua Bay, BP ...... 3740/17747 Raurimu, TO ...... 3907/17524 Whangarei, ND...... 3543/17419 Redhill, AK ...... 3704/17459 Whinray Scenic Reserve, BP ...... 3814/17733 Rimutaka Forest [=Rimutaka Range], WN ...... Woodhill, AK ...... 3644/17425 ...... 4119/17500 Rimutaka Hill, WN ...... 4107/17513 Ruakaka, ND ...... 3554/17427 Ruatoria State Forest, BP ...... 3747/17812 Fauna of New Zealand 53 93

ADDENDUM Hakaharpalus cavelli (Broun, 1893) E new combination See Figures opposite. Tachys (?) cavelli Broun, 1893: 1400. Holotype: female (BMNH) labelled “Type (circular red-bordered label; typed) / [female symbol] (hand-written) / 2442. (hand- written) / New Zeal. Broun Coll. Britt. Mus. 1922—482. (typed) / Capleston Westland. (hand-written) / Tachys cavelli. (hand-written) / Hakaharpalus cavelli (Broun, 1893) new combination Larochelle & Larivière, 2005.” Fair condition. Description. Body length: 4.9 mm. Slightly convex. Brown- ish; margins and sutures of elytra, as well as antennae, palpi, and legs pale yellowish. Generally glabrous. Elytral intervals finely punctate; interneurs shallow (weakly im- pressed), incomplete basally. Microsculpture absent on head, pronotum, and elytra. Head, pronotum, and elytra shiny; dorsal surface without metallic lustre. Head. Mod- erately large, narrower across eyes than pronotal apex; excavated anteriorly, slightly convex posteriorly. Eyes strongly reduced, slightly convex, consisting of obliterated facets, narrowly separated from buccal fissure ventrally (by about 1× maximum width of antennal scape). Antennal scape about 3× longer than its maximum width. Thorax. Pronotum (Fig. opposite) subcordate, widest before mid- dle; sides converging toward base, sinuate; base straight; apex almost straight behind anterior angles; lateral depres- sions absent; anterior angles strongly developed, acute; posterior angles moderately developed, obtuse; basal foveae shallow, ill-defined; punctuation feebly developed. Metepisterna wider than long. Elytra. Widest about mid- dle. Shoulders feebly developed, rounded, without a tooth. Subapical sinuations feeble. Sutural apices rounded. Scutellar striole absent. Interneurs shallow (weakly im- pressed), impunctate, incomplete basally. Intervals finely punctate, flat. Interval 3 without setiferous puncture be- hind middle. Aedeagus. No male known. Material examined. Holotype (BMNH). Geographic distribution. South Island: BR–Capleston. Ecology. Lowland. A wet beech forest. Biology. Unknown. Dispersal power. Elytra fused along suture. Subapterous. Moderate runner (after leg morphology). Remarks. The tribal and generic placements of this spe- cies have remained a mystery for a long time. Examination of the holotype of Tachys (?) cavelli has shown that this taxon belongs neither to the Bembidiini nor to the Zolini as previously thought by a number of carabid workers. Su- perficially, it may resemble members of the Trechinae gen- era Oopterus (Zolini) or Molopsida (Tropopterini) but the Hakaharpalus cavelli (Broun, 1893). Upper: absence of setiferous puncture in the mandibular scrobe holotype, BMNH (scale line =1 mm.); Lower: refers it to the Harpalinae. The authors found this taxon to pronotum (Photographs: M.-C. Larivière) Holotype be congeneric with other species of the newly described photographed with the permission of the Natural genus Hakaharpalus, the characteristic features of which History Museum, London. are given on p. 54. 94 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Fig. 1 Schematic dorsal view of carabid. Fauna of New Zealand 53 95

Fig. 2 Schematic ventral view of carabid. 96 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Fig. 3, 4 (3) Schematic view of pronotum. (4) Schematic view of right elytron. Fauna of New Zealand 53 97

Fig. 5–18 (5-8) Schematic view of microsculpture: (5) granulate; (6) isodiametric; (7) moderately transverse; (8) very transverse. (9-11) Pubescence on anterior margin of penultimate segment of labial palpi: (9) plurisetose; (10) trisetose; (11) bisetose. (12-13) Ventral view of male pro- or mesotarsi: (12) spongily pubescent; (13) biseriately pubescent. (14-18) Medial tooth of mentum: (14) moderately long; (15) longer than lateral lobes; (16) as long as lateral lobes; (17) very short; (18) lacking. 98 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Fig. 19–26 (19-21) Eye and buccal fissure, lateral view: (19) widely separated; (20) narrowly separated; (21) touching (eye reaching buccal fissure). (22-24) Transverse suture between mentum and submentum: (22) complete; (23) incomplete laterally; (24) lacking. (25-26) Membranous laminae of pro- and mesotarsomere 4: (25) present; (26) lacking. Fauna of New Zealand 53 99

Fig. 27–31 (27-28) Pubescence of venter: (27) paired ambulatory setae only; (28) paired ambulatory setae, numerous short setae, and male setiferous fovea. (29-30) Subapical sinuation of right elytron: (29) weak; (30) strong. (31) Ligula as long as paraglossae. 100 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Fig. 32-84 Aedeagus. Lateral view, except when indicated otherwise. Fauna of New Zealand 53 101 102 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 103 104 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 105 106 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 107

Fig. 85-113 Habitus drawings of representatives of Harpalini (Illustrator: D. W. Helmore). Scale lines are 1 mm. 108 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 109 110 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 111 112 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 113 114 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 115

Fig. 114-169 Colour photographs of Harpalini pronota. (Photographer: M.-C. Larivière). 116 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 117 118 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 119 120 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 121 122 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 123 124 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 125

Fig. 170–225 Colour photographs of Harpalini. (Photographer: B. E. Rhode, except Fig. 187 M.-C. Larivière). Scale lines are 1 mm. 126 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 127 128 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 129 130 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 131 132 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 133 134 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 135 136 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 137 138 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 139 140 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Map 1 The New Zealand subregion. Fauna of New Zealand 53 141

Map 2 Area codes and collecting localities from mainland New Zealand: North Island. 142 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Map 3 Area codes and collecting localities from mainland New Zealand: South Island and Stewart Island. Fauna of New Zealand 53 143

Map 4 Total number of known taxa by areas. 144 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Map 5 Number of known New Zealand endemics by areas. Fauna of New Zealand 53 145

Map 6 Number of native taxa known to be restricted to single areas. 146 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

Map 7 Number of known adventive taxa by areas.

Species distribution maps (pp. 147-153). Presented in same order as taxa in body of text. Area boundaries follow area codes of Crosby et al. (1976, 1998). Fauna of New Zealand 53 147 148 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 149 150 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 151 152 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) Fauna of New Zealand 53 153 154 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

TAXONOMIC INDEX antarcticus (Laporte de clunieae Larochelle & Castelnau), Hypharpax 14, Larivière, Tuiharpalus 17, 18, This index covers the nominal 16, 18, 19, 38, 101f, 116p, 19, 26, 46, 47, 48, 49, 102f, taxa mentioned in the text, regard- 127p, 148m 118p, 130p, 149m less of their current status in tax- atratus Broun, Parabaris 18, coxii Laporte de Castelnau, onomy. In the case of synonyms, 19, 42, 43, 44, 101f, 109f, Harpalus 39 the combinations of generic and 117p, 128p, 148m crosbyi Larochelle & Larivière, specific names listed are those atriceps (Macleay), Tuiharpalus 16, 17, 18, 19, originally published by authors, Lecanomerus 18, 19, 20, 60, 47, 102f, 109f, 118p, 129p, and may differ from combinations 61, 62, 77, 103f, 120p, 133p, 149m implicit in current usage. Taxa in 150m crypticus Moore, Haplanister bold indicate valid taxa. Page atriceps Macleay, Trechus 62 16, 17, 18, 19, 78, 105f, 113f, numbers in bold indicate main atroviridis Macleay, Harpalus 37 123p, 138p, 152m catalogue entries. The letter “f” australasiae Dejean, Harpalus after a page indicates a figure. 18, 19, 20, 51, 53, 102f, 111f, davidsoni Larochelle & The letter “m” indicates a distri- 119p, 131p, 149m Larivière, Hakaharpalus 17, bution map. The letter “p” indi- australis Dejean, Harpalus 39 18, 19, 55, 56, 103f, 119p, cates a photograph. australis (Dejean), Hypharpax 132p, 149m 18, 19, 20, 26, 38, 39, 101f, Diaphoromerus Chaudoir 41 108f, 116p, 127p, 148m dingo Laporte de Castelnau, abstrusus Bates, Hypharpax 39 Harpalus 73 adelaideae Laporte de barrattae Larochelle & Castelnau, Harpalus 37 Larivière, Kupeharpalus 18, Egadroma Motschulsky 16, 20, aeneonitens Macleay, Harpalus 19, 58, 59, 63, 103f, 111f, 25, 72 37 120p, 133p, 150m embersoni Larochelle & aeneus Fabricius, Carabus 51 belli Larochelle & Larivière, Larivière, Kupeharpalus 17, affinis Schrank, Carabus 51 Allocinopus 17, 18, 19, 28, 18, 19, 58, 59, 63, 103f, 120p, affinis (Schrank), Harpalus 30, 31, 100f, 115p, 125p, 133p, 150m 18,19, 51, 102f, 110f, 119p, 147m Eurytrichus LeConte 41 131p, 149m bicolor Moore, Euthenarus 18, Euthenarus Bates 15, 16, 20, Allocinopus Broun 16, 25, 26, 19, 20, 62, 74, 76, 105f, 123p, 25, 72, 73 27 137p, 152m Amblystus Motschulsky 51 binotatus (Fabricius), fallax Broun, Lecanomerus 63 angustatus Macleay, Harpalus Anisodactylus 16, 18, 19, 34, flavocinctus Blackburn, 37 101f, 107f, 116p, 126p, 147m Lecanomerus 67 angustulus Broun, Allocinopus bousqueti Larochelle & fuliginosus Broun, 18, 19, 26, 28, 29, 37, 100f, Larivière, Allocinopus 17, 18, Lecanomerus 64 115p, 125p, 147m 19, 28, 31, 100f, 115p, 125p, fulvescens Bates, Triplosarus Anisodactyli 15 147m 45, 46 Anisodactylina 15, 18, 19, 20, brevicollis Bates, Euthenarus 25, 26, 27 16, 18, 19, 74, 76, 105f, 122p, Gaioxenus Broun 16, 25, 26, Anisodactylus Dejean 16, 25, 137p, 152m 27, 35 26, 27, 34 gayndahensis Macleay, Anisodactylus (Anisodactylus) castaneus Broun, Allocinopus Harpalus 37 34 28 Gnathaphanus Macleay 16, 20, Anisotarsus Chaudoir 41 cavelli (Broun), Hakaharpalus 25, 26, 27, 36 anomalus Bates, Syllectus 16, 17, 55, 57, 93p gouleti Larochelle & Larivière, 18, 19, 69, 104f, 112f, 122p, cavelli Broun, Tachys 57, 93 Syllectus 17, 18, 19, 69, 71, 136p, 151m cerberus Britton, Pholeodytes 104f, 122p, 136p, 151m antarcticus Laporte de 17, 18, 19, 80, 81, 82, 106f, gourlayi Britton, Parabaris 46, Castelnau, Harpalus 38 124p, 139p, 152m 48 Fauna of New Zealand 53 155 gourlayi (Britton), Tuiharpalus labralis Broun, Lecanomerus novaezelandiae Laporte de 16, 17, 18, 19, 46, 47, 48, 67 Castelnau, Harpalus 14, 45 102f, 110f, 118p, 129p, 149m latimanus Bates, novaezelandiae (Laporte de Lecanomerus 16, 18, 19, 61, Castelnau), Triplosarus 16, Hakaharpalus Larochelle & 64, 104f, 121p, 134p, 150m 17, 18, 19, 45, 102f, 109f, Larivière 15, 16, 17, 25, 54, latitarsis Broun, Allocinopus 117p, 129p, 148m 93 16, 18, 19, 28, 32, 100f, 115p, nunni Larochelle & Larivière, hallae Larochelle & Larivière, 126p, 147m Pholeodytes 17, 18, 19, 80, Tuiharpalus 18, 19, 26, 46, Lecanomerus Chaudoir 15, 16, 82, 106f, 124p, 139p, 152m 47, 49, 102f, 110f, 118p, 130p, 20, 25, 54, 58, 60, 63 149m lesagei Larochelle & Larivière, obesulus Bates, Lecanomerus Haplanister Moore 14, 16, 25, Parabaris 17, 18, 19, 42, 43, 18, 19, 61, 63, 65, 104f, 121p, 72, 77 101f, 117p, 128p, 148m 134p, 151m Harpalina 15, 18, 19, 20, 25, 50 occidentalis Sloane, Harpalini 24, 25 maddisoni Larochelle & Lecanomerus 67 Harpalus Latreille 16, 20, 25, Larivière, Hakaharpalus 17, ocularius Broun, Allocinopus 33 51 18, 19, 55, 56, 57, 103f, 119p, Odontagonum Darlington 60 Harpalus (Harpalus) 51 132p, 150m helmorei Larochelle & magnus Britton, Syllectus 16, Pachauchenius Macleay 36 Larivière, Pholeodytes 17, 18, 19, 69, 70, 104f, 122p, pallipes Broun, Lecanomerus 18, 19, 80, 83, 124p, 139p, 136p, 151m 64 152m Maoriharpalus Larochelle & palmai Larochelle & Larivière, hoarei Larochelle & Larivière, Larivière 16, 17, 25, 26, 27, Pholeodytes 17, 18, 19, 80, Parabaris 17, 18, 19, 42, 43, 40 81, 106f, 124p, 138p, 153m 44, 101f, 109f, 117p, 129p, marginatus Sharp, Parabaris Broun 14, 16, 25, 27, 148m Lecanomerus 65 42, 46 Hypharpax Macleay 16, 20, 25, marginicollis Laporte de paroensis Laporte de 26, 27, 37, 38 Castelnau, Harpalus 37 Castelnau, Harpalus 36 marrisi Larochelle & Larivière, parvus Chaudoir, Hypharpax 39 incertus Broun, Lecanomerus Lecanomerus 17, 18, 19, 60, patricki Larochelle & Larivière, 64 61, 66, 104f, 112f, 121p, 135p, Hakaharpalus 17, 18, 19, 55, inornatus Germar, Harpalus 39 151m 57, 103f, 111f, 119p, 132p, insidiosus Chaudoir, mastersii Macleay, Acupalpus 150m Lecanomerus 67 67 Pelmatellina 15, 18, 19, 20, 25, insignitus Broun, melbournensis (Laporte de 54 Lecanomerus 18, 19, 60, 61, Castelnau), Gnathaphanus Pholeodytes Britton 14, 15, 16, 63, 103f, 112f, 120p, 134p, 18, 19, 20, 36, 101f, 108f, 17, 25, 69, 71, 72, 80 150m 116p, 127p, 148m picea (Guérin-Méneville), insularis Bates, Mirosarus 37 melbournensis Laporte de Egadroma 18, 19, 20, 73, Castelnau, Harpalus 36 105f, 113f, 122p, 137p, 151m javanus Jedli…ka, Acupalpus 62 Mirosarus Bates 36 piceus Guérin-Méneville, johnsi Larochelle & Larivière, moorei Larochelle & Larivière, Acupalpus 73 Kupeharpalus 17, 18, 19, 58, Tuiharpalus 17, 18, 19, 46, pilipalpis Broun, Gaioxenus 18, 59, 103f, 111f, 120p, 133p, 47, 50, 102f, 110f, 118p, 130p, 19, 35, 101f, 107f, 116p, 127p, 150m 149m 147m planipennis Macleay, Harpalus Kiwiharpalus Larochelle & nitidus Blackburn, 37 Larivière 15, 16, 17, 25, 72, Lecanomerus 67 politus Macleay, Stenolophus 73 79 Notiobia Perty 20, 25, 26, 27, Polpochili 15 Kupeharpalus Larochelle & 41 promptus (Erichson), Larivière 15, 16, 25, 54, 57, Notiobia (Anisotarsus) 41 Euthenarus 18, 19, 20, 74, 60 Notiobii 15 77, 105f, 123p, 138p, 152m 156 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae) promptus Erichson, Harpalus sharpi Csiki, Nemaglossa 65 towsendi Britton, Pholeodytes 77 smithi Broun, Allocinopus 18, 18, 19, 80, 82, 106f, 114f, puncticollis Bates, Euthenarus 19, 26, 28, 100f, 107f, 115p, 124p, 139p, 153m 16, 18, 19, 74, 75, 105f, 113f, 125p, 147m Triplosarus Bates 25, 27, 45 122p, 137p, 152m spelaeus Britton, Syllectus 70, Tuiharpalus Larochelle & 71 Larivière 16, 25, 27, 46 quadricollis Chaudoir, Stenolophi 15 Diaphoromerus 41 Stenolophina 15, 18, 19, 25, 72 verticalis Erichson, Harpalus 67 quadricollis (Chaudoir), stenopus Broun, Lecanomerus verticalis (Erichson), Notiobia 18, 19, 20, 41, 108f, 67 Lecanomerus 18, 19, 20, 60, 117p, 128p, 148m Stilbolidus Casey 41 61, 67, 68, 104f, 121p, 135p, sutherlandi Larochelle & 151m rhodeae Larochelle & Larivière, Maoriharpalus 16, vestigialis Erichson, Harpalus Larivière, Hakaharpalus 17, 17, 18, 19, 40, 101f, 108f, 67 18, 19, 55, 57, 103f, 120p, 117p, 128p, 148m vestigialis (Erichson), 132p, 150m Syllectus Bates 14, 15, 16, 17, Lecanomerus 18, 19, 20, 60, 25, 54, 68, 71, 80 61, 67, 104f, 112f, 121p, 135p, Sagraemerus Redtenbacher 37 151m sculpticollis Broun, tardus Panzer, Carabus 52 Allocinopus 16, 18, 19, 28, tardus (Panzer), Harpalus 18, wardi Larochelle & Larivière, 33, 100f, 107f, 116p, 126p, 19, 51, 52, 102f, 119p, 131p, Allocinopus 17, 18, 19, 28, 147m 149m 31, 100f, 115p, 126p, 147m sexualis Fauvel, Stenolophus Thenarotes Bates 60 73 townsendi Larochelle & 2notatus Fabricius, Carabus 34 sharpi (Csiki), Lecanomerus Larivière, Kiwiharpalus 16, 16, 18, 19, 60, 61, 65, 104f, 17, 18, 19, 79, 113f, 123p, 121p, 134p, 151m 138p, 152m Fauna of New Zealand 53 157

TITLES IN PRINT / PUNA TAITARA TAA

1 Terebrantia (Insecta: Thysanoptera) • Laurence A. Mound & Annette K. Walker ISBN 0-477-06687-9 • 23 Dec 1982 • 120 pp...... $29.95 2 Osoriinae (Insecta: Coleoptera: Staphylinidae) • H. Pauline McColl ISBN 0-477-06688-7 • 23 Dec 1982 • 96 pp...... $18.60 3 Anthribidae (Insecta: Coleoptera) • B.A. Holloway ISBN 0-477-06703-4 • 23 Dec 1982 • 272 pp...... $41.00 4 Eriophyoidea except Eriophyinae (Arachnida: Acari) • D.C.M. Manson ISBN 0-477-06745-X • 12 Nov 1984 • 144 pp...... $29.95 5 Eriophyinae (Arachnida: Acari: Eriophyoidea) • D.C.M. Manson ISBN 0-477-06746-8 • 14 Nov 1984 • 128 pp...... $29.95 6 Hydraenidae (Insecta: Coleoptera) • R.G. Ordish ISBN 0-477-06747-6 • 12 Nov 1984 • 64 pp...... $18.60 7 Cryptostigmata (Arachnida: Acari) – a concise review • M. Luxton ISBN 0-477-06762-X • 8 Dec 1985 • 112 pp...... $29.95 8 Calliphoridae (Insecta: Diptera) • James P. Dear ISBN 0-477-06764-6 • 24 Feb 1986 • 88 pp...... $18.60 9 Protura (Insecta) • S.L. Tuxen ISBN 0-477-06765-4 • 24 Feb 1986 • 52 pp...... $18.60 10 Tubulifera (Insecta: Thysanoptera) • Laurence A. Mound & Annette K. Walker ISBN 0-477-06784-0 • 22 Sep 1986 • 144 pp...... $34.65 11 Pseudococcidae (Insecta: Hemiptera) • J.M. Cox ISBN 0-477-06791-3 • 7 Apr 1987 • 232 pp...... $49.95 12 Pompilidae (Insecta: Hymenoptera) • A.C. Harris ISBN 0-477-02501-3 • 13 Nov 1987 • 160 pp...... $39.95 13 Encyrtidae (Insecta: Hymenoptera) • J.S. Noyes ISBN 0-477-02517-X • 9 May 1988 • 192 pp...... $44.95 14 Lepidoptera – annotated catalogue, and keys to family-group taxa J. S. Dugdale • ISBN 0-477-02518-8 • 23 Sep 1988 • 264 pp...... $49.95 15 Ambositrinae (Insecta: Hymenoptera: Diapriidae) • I.D. Naumann ISBN 0-477-02535-8 • 30 Dec 1988 • 168 pp...... $39.95 16 Nepticulidae (Insecta: Lepidoptera) • Hans Donner & Christopher Wilkinson ISBN 0-477-02538-2 • 28 Apr 1989 • 92 pp...... $22.95 17 Mymaridae (Insecta: Hymenoptera) – introduction, and review of genera J.S. Noyes & E.W. Valentine • ISBN 0-477-02542-0 • 28 Apr 1989 • 100 pp...... $24.95 18 Chalcidoidea (Insecta: Hymenoptera) – introduction, and review of genera in smaller families J.S. Noyes & E.W. Valentine • ISBN 0-477-02545-5 • 2 Aug 1989 • 96 pp...... $24.95 19 Mantodea (Insecta), with a review of aspects of functional morphology and biology • G.W. Ramsay • ISBN 0-477-02581-1 • 13 Jun 1990 • 96 pp...... $24.95 20 Bibionidae (Insecta: Diptera) • Roy A. Harrison ISBN 0-477-02595-1 • 13 Nov 1990 • 28 pp...... $14.95 21 Margarodidae (Insecta: Hemiptera) • C.F. Morales ISBN 0-477-02607-9 • 27 May 1991 • 124 pp...... $34.95 22 Notonemouridae (Insecta: Plecoptera) • I.D. McLellan ISBN 0-477-02518-8 • 27 May 1991 • 64 pp...... $24.95 23 Sciapodinae, Medeterinae (Insecta: Diptera) with a generic review of the Dolichopodidae • D.J. Bickel • ISBN 0-477-02627-3 • 13 Jan 1992 • 74 pp...... $27.95 24 Therevidae (Insecta: Diptera) • L. Lyneborg ISBN 0-477-02632-X • 4 Mar 1992 • 140 pp...... $34.95 25 Cercopidae (Insecta: Homoptera) • K.G.A. Hamilton & C.F. Morales ISBN 0-477-02636-2 • 25 May 1992 • 40 pp...... $17.95 158 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

26 Tenebrionidae (Insecta: Coleoptera): catalogue of types and keys to taxa J.C. Watt • ISBN 0-477-02639-7 • 13 Jul1992 • 70 pp...... $27.95 27 Antarctoperlinae (Insecta: Plecoptera) • I.D. McLellan ISBN 0-477-01644-8 • 18 Feb 1993 • 70 pp...... $27.95 28 Larvae of Curculionoidea (Insecta: Coleoptera): a systematic overview Brenda M. May • ISBN 0-478-04505-0 • 14 Jun 1993 • 226 pp...... $55.00 29 Cryptorhynchinae (Insecta: Coleoptera: Curculionidae) C.H.C. Lyal • ISBN 0-478-04518-2 • 2 Dec 1993 • 308 pp...... $65.00 30 Hepialidae (Insecta: Lepidoptera) • J.S. Dugdale ISBN 0-478-04524-7 • 1 Mar 1994 • 164 pp...... $42.50 31 Talitridae (Crustacea: Amphipoda) • K.W. Duncan ISBN 0-478-04533-6 • 7 Oct 1994 • 128 pp...... $36.00 32 Sphecidae (Insecta: Hymenoptera) • A.C. Harris ISBN 0-478-04534-4 • 7 Oct 1994 • 112 pp...... $33.50 33 Moranilini (Insecta: Hymenoptera) • J.A. Berry ISBN 0-478-04538-7 • 8 May 1995 • 82 pp...... $29.95 34 Anthicidae (Insecta: Coleoptera) • F.G. Werner & D.S. Chandler ISBN 0-478-04547-6 • 21 Jun 1995 • 64 pp...... $26.50 35 Cydnidae, Acanthosomatidae, and Pentatomidae (Insecta: Heteroptera): systematics, geographical distribution, and bioecology • M.-C. Larivière ISBN 0-478-09301-2 • 23 Nov 1995 • 112 pp...... $42.50 36 Leptophlebiidae (Insecta: Ephemeroptera) • D.R. Towns & W.L. Peters ISBN 0-478-09303-9 • 19 Aug 1996 • 144 pp...... $39.50 37 Coleoptera: family-group review and keys to identification • J. Klimaszewski & J.C. Watt • ISBN 0-478-09312-8 • 13 Aug 1997 • 199 pp...... $49.50 38 Naturalised terrestrial Stylommatophora (Mollusca: Gastropoda) • G.M. Barker ISBN 0-478-09322-5 • 25 Jan 1999 • 253 pp...... $72.50 39 Molytini (Insecta: Coleoptera: Curculionidae: Molytinae) • R.C. Craw ISBN 0-478-09325-X • 4 Feb1999 • 68 pp...... $29.50 40 Cixiidae (Insecta: Hemiptera: Auchenorrhyncha) • M.-C. Larivière ISBN 0-478-09334-9 • 12 Nov 1999 • 93 pp...... $37.50 41 Coccidae (Insecta: Hemiptera: Coccoidea) • C. J. Hodgson & R. C. Henderson ISBN 0-478-09335-7 • 23 Feb 2000 • 264 pp...... $72.50 42 Aphodiinae (Insecta: Coleoptera: Scarabaeidae) • Z. T. Stebnicka ISBN 0-478-09341-1 • 15 Jun 2001 • 64 pp...... $29.50 43 Carabidae (Insecta: Coleoptera): catalogue • A. Larochelle & M.-C. Larivière ISBN 0-478-09342-X • 15 Jun 2001 • 285 pp...... $72.50 44 Lycosidae (Arachnida: Araneae) • C. J. Vink ISBN 0-478-09347-0 • 23 Dec 2002 • 94 pp...... $37.50 45 Nemonychidae, Belidae, Brentidae (Insecta : Coleoptera : Curculionoidea) • G. Kuschel ISBN 0-478-09348-9 • 28 Apr 2003 • 100 pp...... $40.00 46 Nesameletidae (Insecta : Ephemeroptera) • T. R. Hitchings & A. H. Staniczek ISBN 0-478-09349-7 • 14 May 2003 • 72 pp...... $32.50 47 Erotylidae (Insecta : Coleoptera: Cucujoidea): phylogeny and review • R. A. B. Leschen ISBN 0-478-09350-0 • 5 June 2003 • 108 pp...... $42.50 48 Scaphidiinae (Insecta : Coleoptera: Staphylinidae) • I. Löbl & R. A. B. Leschen ISBN 0-478-09353-5 • 18 Nov 2003 • 94 pp...... $37.50 49 Lithinini (Insecta : Lepidoptera: Geometridae: Ennominae) • J. D. Weintraub & M. J. Scoble ISBN 0-478-09357-8 • 29 Apr 2004 • 48 pp...... $24.50 50 Heteroptera (Insecta : Hemiptera): catalogue • M.-C. Larivière & A. Larochelle ISBN 0-478-09358-6 • 14 May 2004 • 330 pp...... $89.00 51 Coccidae (Insecta : Hemiptera: Coccoidea): adult males, pupae and prepupae of indigenous species C. J. Hodgson & R. C. Henderson • ISBN 0-478-09360-8 • 22 June 2004 • 228 pp...... $65.00 52 Raphignathoidea (Acari: Prostigmata) • Qing-Hai Fan & Zhi-Qiang Zhang ISBN 0-478-09360-8 • May 2005 • 400 pp...... $89.00 53 Harpalini (Insecta : Coleoptera: Carabidae: Harpalinae) • A. Larochelle & M.-C. Larivière ISBN 0-478-09369-1 • May 2005 • 160 pp...... $55.00 Visit the Manaaki Whenua Press Website at http://www.mwpress.co.nz/ for further information, and to gain access to on-line extracts from these publications. Fauna of New Zealand 53 159

Taxonomic groups covered in the Hymenoptera Chalcidoidea: introduction, and review of smaller families Fauna of New Zealand series (J.S. Noyes & E.W. Valentine, FNZ 18, 1989) Diapriidae: Ambositrinae (I.D. Naumann, FNZ 15, 1988) Insecta Encyrtidae (J.S. Noyes, FNZ 13, 1988) Coleoptera Mymaridae (J.S. Noyes & E.W. Valentine, FNZ 17, 1989) Family-group review and keys to identification (J. Pompilidae (A.C. Harris, FNZ 12, 1987) Klimaszewski & J.C. Watt, FNZ 37, 1997) Pteromalidae: Eunotinae: Moranilini (J.A. Berry, FNZ 33, Anthribidae (B.A. Holloway, FNZ 3, 1982) 1995) Anthicidae (F.G. Werner & D.S. Chandler, FNZ 34, 1995) Sphecidae (A.C. Harris, FNZ 32, 1994) Carabidae: catalogue (A. Larochelle & M.-C. Larivière, FNZ 43, 2001) Lepidoptera Carabidae: Harpalinae: Harpalini (A. Larochelle & M.-C. Annotated catalogue, and keys to family-group taxa (J. S. Larivière, FNZ 53, 2005) Dugdale, FNZ 14, 1988) Curculionidae: Cryptorhynchinae (C.H.C. Lyal, FNZ 29, Geometridae: Ennominae: Lithinini (Jason D. Weintraub & 1993) Malcolm J. Scoble, FNZ 49, 2004) Curculionidae: Molytinae: Molytini (R. C. Craw, FNZ 39, Hepialidae (J.S. Dugdale, FNZ 30, 1994) 1999) Nepticulidae (Hans Donner & Christopher Wilkinson, FNZ Curculionoidea: Nemonychidae, Belidae, Brentidae (G. 16, 1989) Kuschel, FNZ 45, 2003) Curculionoidea larvae: a systematic overview (Brenda M. Mantodea, with a review of aspects of functional mor- May, FNZ 28, 1993) phology and biology (G.W. Ramsay, FNZ 19, 1990) Erotylidae: phylogeny and review (Richard A. B. Leschen, FNZ 47, 2003) Plecoptera Hydraenidae (R.G. Ordish, FNZ 6, 1984) Antarctoperlinae (I.D. McLellan, FNZ 27, 1993) Scarabaeidae: Aphodiinae (Z. T. Stebnicka, FNZ 42, 2001) Notonemouridae (I.D. McLellan, FNZ 22, 1991) Staphylinidae: Osoriinae (H. Pauline McColl, FNZ 2, 1982) Staphylinidae: Scaphidiinae (I. Löbl & Richard A. B. Protura (S.L. Tuxen, FNZ 9, 1986) Leschen, FNZ 48, 2003) Tenebrionidae: catalogue of types and keys to taxa (J.C. Thysanoptera Watt, FNZ 26, 1992) Terebrantia (Laurence A. Mound & Annette K. Walker, FNZ 1, 1982) Diptera Tubulifera (Laurence A. Mound & Annette K. Walker, FNZ Bibionidae (Roy A. Harrison, FNZ 20, 1990) 10, 1986) Calliphoridae (James P. Dear, FNZ 8, 1986) Dolichopodidae: Sciapodinae, Medeterinae with a generic Arachnida review (D.J. Bickel, FNZ 23, 1992) Acari Therevidae (L. Lyneborg, FNZ 24, 1992) Cryptostigmata – a concise review (M. Luxton, FNZ 7, 1985) Ephemeroptera Eriophyoidea except Eriophyinae (D.C.M. Manson, FNZ 4, Leptophlebiidae (D.R. Towns & W.L. Peters, FNZ 36, 1996) 1984) Nesameletidae (Terry R. Hitchings & Arnold H. Staniczek, Eriophyinae (D.C.M. Manson, FNZ 5, 1984) FNZ 46, 2003) Raphignathoidea (Qing-Hai Fan & Zhi-Qiang Zhang, FNZ 52, 2005) Hemiptera Cercopidae (K.G.A. Hamilton & C.F. Morales, FNZ 25, Araneae 1992) Lycosidae (C. J. Vink, FNZ 44, 2002) Cixiidae (M.-C. Larivière, FNZ 40, 1999) Coccidae (C. J. Hodgson & R. C. Henderson, FNZ 41, Crustacea 2000); adult males, pupae and prepupae of indig- Amphipoda enous species (C. J. Hodgson & R. C. Henderson, Talitridae (K.W. Duncan, FNZ 31, 1994) FNZ 51, 2004) Cydnidae, Acanthosomatidae, and Pentatomidae (M.-C. Mollusca Larivière, FNZ 35, 1995) Gastropoda Heteroptera: catalogue (M.-C. Larivière & A. Larochelle, Naturalised terrestrial Stylommatophora (G.M. Barker, FNZ FNZ 50, 2004) 38, 1999) Margarodidae (C.F. Morales, FNZ 21, 1991) Pseudococcidae (J.M. Cox, FNZ 11, 1987) 160 Larochelle & Larivière (2005): Harpalini (Insecta: Coleoptera: Carabidae)

NOTICES NGA PANUI

This series of refereed publications has been established to Kua whakatãria t‘nei huinga pukapuka hei whakahauhau i encourage those with expert knowledge to publish concise ng~ tohunga whai m~tauranga kia whakaputa i ng~ kÇrero yet comprehensive accounts of elements in the New poto, engari he whaikiko tonu, e p~ ana ki ng~ aitanga Zealand fauna. The series is professional in its conception pepeke o Aotearoa. He tÇtika tonu te ~hua o ng~ tuhituhi, and presentation, yet every effort is made to provide engari ko te tino wh~inga, kia m~rama te marea ki ng~ tohu resources for identification and information that are tautuhi o ia ng~rara, o ia ng~rara, me te roanga atu o ng~ accessible to the non-specialist. kÇrero mÇ t‘n~, mÇ t‘n~.

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