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A CA E ESEA C E ESE A I ES
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U I E SI IES E ESE A I E r .M. E b r n p rt nt f Ent l & An l E l .O. x 84, n ln Un v r t , l nd
MUSEUMS E ESE A I E Mr . . l t r l Env r n nt p rt nt M f l nd p n r .O. x 46 , W ll n t n, l nd
O E SEAS E ESE A I E r . . r n CSI O v n f Ent l G. .O. x 00, C nb rr C t A.C. . 260 , A tr l
SE IES E I O AU A O EW EA A Mr C v l nd r r h M nt Alb rt r h C ntr r v t 2 0, A l nd, l nd Fauna of New Zealand K t A t n Ρ ρ A t r
Number / Nama 37
Coleoptera: family-group review and keys to identification
. Kl z 1 and .C. W tt 2 with illustrations by D.W. Helmore
Landcare Research Mount Albert Research Centre Private Bag 92170, Auckland, New Zealand
1 Present address: BC Research Inc. 3650 Wesbrook Mall, Vancouver British Columbia, Canada V6S 2L2
2 Research Associate
Manaaki Whenua PRESS Lincoln, Canterbury, New Zealand 1997
1 C p r ht © nd r r h l nd td 1997
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PubIication of the Fauna of New Zealand r s supported by the Foundation for Research, Science and TechnoIogy under contract nos C09308 and C09617. O U A SUMMA Y E W AKA O O GA ΜΑ E MA EA
Cl In t Ord r C l pt r
Families of beetles
tl r th l r t rd r f r n pr n v r 35 d r b d p n th rld f n h p - r l f d n 3 n r It h b n t - t d th t th r r r p f b tl th n th r r Ill tr t n Wh h : M r nt nd p r n f v l r pl nt r f n nd 9 t n th r h lp t l f b tl nt f l (Ill tr t r / K - r l p h h K r t W l n In l nd t l t 51 n t v p f b tl h v b n r n d A f rth r 3 3 p r th r lf- O n ā pūt r r p t t pūt r r t t n ntr d d r h v b n d l b r t l br ht n f r x- n t t 35 n ā r r t t pl f r th b l l ntr l f n x d h t t h p t n t E n h n ēn b tl b l n t 1 9 n r n f l ēt h p n n āh 3 n E t h r p r n th v l r pl nt fl r n l d b t 5 ēt h h h n ā r r tēnā n ā p nd th t rr tr l v rt br t f n b t 35 p - t p h n ā ōp r h tū rān e 90 te h t l n b r f b tl p n l nd h r n n ā r r n ā r - r h h n t nd bt dl h h h r — n r nd r b d t t 51 n ā r r A t r h f r t E r p n ll t n f l nd b tl t t t tēn h n Arā nō ēt h r d b ph n nd n l S l nd r d r n momo e 343 nei tau noa āt rān pērā C pt n C f r t v n 17 9 nd th f r t ērā h nōh h p n ā t r kino. E noho f r l d r pt n r b h nn br n 1775 katoa n ēn ēt h p n n 1 9 ēt h h r n f l nd b tl nt ll n- hān K t h r t tēn t n ēt h t n n b t pp r ntl t l t h d d v l p d n n r r p 5 n ā t p h ā 35 n h n nt l nd f nt n nt l r n h h b n- n ā r r h n h h n h t rā n Kā r r n l l t d n th l t M z r h rl - e kore he nui noa n ā r r A t r tērā t n n b tl f l fr l nd f n n - h n r n n nā h rā nō ēt h ā r nō pl t fr nt n d t n fr th Cr t āt h h h āt t t h r f t r r r p n bl f r th p t n nd K ph n rā n l S l nd r n ā ā hā d v r t f th pr nt-d b tl f n f l nd t t h t h r t āt h h n ā r r (1 h r n f f r f n n th n nt th- A t r rā h r t r r n t t h rn p r nt n nt n n G nd n ( h t t d Hē K A t r t t 17 9 ā n ā t t h n ll n r f r ph l t n f l nd whaitake tuatahi, n l Johanns Fabricius i te tau 1775. h h h r lt d n 9 p r nt nd (l l Kā r āt ōh t n t pūt n n ā r n f p nd h h nd l t n l v l r r A t r n r t ōr r rā ēt h (3 Ch n n l t l nd r nd l nd rf d t t tēt h h n r h tūā r r M i te f t r h l t n nt n b ld n nd v l n āh n h t n t ā M z n h r t v t ( h b n f l nd th r n l tēn h n r h t aba, ka motuhake ēt h t nd pl nt h h h v d n t d t l h r whenua. Ko te parawae kurikuri tino tawhito o Aotearoa
( nt n d v rl f ( r h r tοn in the world. The flora and fauna have been greatly modi- ōh t n tēt h āh n t p h fied in the last 1 years by humans, who brought devas- t ā Cr t tation to native forest and introduced exotic animals with E hā n ā āh t n h t h t āh t n destructive influence (e.g., deer, rats, possum, goats). n ā r r A t r ēn r (1 K t Contemporary New Zealand beetles are thus a compos- pūt n ēt h t h n r h tūā r r ite of ancient, variously changed lineages, elements intro- ī n G nd n ( K t āh r n t duced by dispersal over short and long distances, and spe- n h t h n A t r n 9 % t cies intentionally and accidentally introduced by humans. n n ā r r A t r nō n t t ā The majority of species are associated with native forest, t ēn ēt h t h n W h which now constitutes only 23 percent of the total land t pērā n t āh n ā p n n (3 Κ t r r ē area. The shrinking forest has profoundly reduced the popu- h r t āh n ā r n t t t t h n lation size of many native species and their potential for t t t t t h n ā n ā āh t n pēn long-term survival. n ā h p p n ā āh t n h n n n ā Our beetles are also vulnerable to introduced organisms: h ū pēn ( Κ t r ēt h their defence mechanisms are often ineffective against the r r h h n t ēt h t r r h t t new arrivals. They may succumb to competition for food ēt h t p n h ātā ēt h t r r n h n t I tēn 1 with introduced wasps, to predation by mammals, and to t h n n h t n t r h n ā t p n ā r r h depletion of food sources by weeds out-competing native A t r t r n t n t t t n t ē h r plants. A number of beetle species are recognised as being h r n ā n h r ā r n n nδ t t h threatened with extinction, and are listed for protection by t r r h n t h t (h t r rā t t t r the Department of Conservation. t p h t t On a positive note, beetle diversity can be unexpectedly Na r r n ā r r A t r nā n h high in relatively small areas of habitat. The beetle fauna t h r ōt ēn āh t n h n n ā of the Auckland suburb of Lynfield has been shown to ā n h r t r hā n ā t ā- exceed 1000 species, of which three-quarters are native. p n nī ā- n p rān h h p t h t p t t rān t t n ā āt h r t t n t h r p r hū rān t t n t Κ t Contributor n Kl z was born in Poland in n n n ā n h n n n ā n āh r 1950. He graduated MSc in 1973 and PhD in 1978 from ā r ā t t n ā h n A t r tēt h the University of Wroclaw, with a specialisation in the 3% n h ā r t r r n ā n h r beetle family Staphylinidae. He was based in Poland until ā r t t h tēnā tēnā 1980, when he took up a postdoctoral research fellowship n ā r r tūt r A t r ā tērā p r in Ottawa, Canada. From 1982 to 1989 he was a research n h pū t t h n associate in the Lyman Entomological Museum at McGill K ōr r nō h ā tāt r r t n h University, working on lacewings (Neuroptera). Jan was t h Id n ā r r p t h n ē h then appointed as a Senior Curator of Coleoptera in the r t rāt t p r t n ā t tērā p pā Transvaal Museum, Pretoria, South Africa. In 1993 he ēn r r p h ērā n h p p t h - returned briefly to Canada before taking up the post of t t t h n ā t p h r t h t — r r Coleopterist with Landcare Research in New Zealand. h n t t h ; n ā r r t n rān n h h tēt h r r h h n t ; r r Contributor Ch rl W tt was born in England but rān n ā t rn t h t ē tā n ā t p n t - moved to New Zealand at an early age. He graduated MSc t t h nō ēn p pār t n n ā (Hons) in zoology from the University of Auckland in 1960. r r In 1965 he graduated DPhil from Oxford University, where Arā ēt h t t t n t r hāhā ā h he worked on the beetle family Tenebrionidae. He then h r ēn p At h t rār n n ā joined the Entomology Division of DSIR, with responsi- r r p āt h āt r h t bility in the Systematics Section for all beetles except the nō t r h t t h r n p rā ēt h āh weevils. Charles was obliged to retire prematurely in 1986 n h r r n ē t n n ā r r n h because of a stroke which left him hemiplegic. He has since n r r E n ā ōr r n t t 1 n ā been an honorary research associate of the invertebrate r r n h t h n t t ā nf ld systematics group, first under DSIR and since 1992 under ā - -r ā tēt h 75% ēn n n Landcare Research. t t E ACE
The idea of producing a manual for identification of New Zealand beetle families originated long ago in the Entomology Division of the former DSIR, yet for some time never quite got off the ground. This changed when, after graduating in 1965 from Oxford University, Charles Watt joined Entomology Division with responsibilities in the Systematics Section for Coleoptera. Over two decades he gathered together representative specimens of many beetle species, compiling their general descriptions, geographic distribution, and data on natural history, and produced a pre- liminary linear key for the identification of our families. His selected examples, which have been meticulously drawn by illustrator Des Helmore, represent most of the families occurring in New Zealand. In October 1985, while working at the Natural History Museum in Paris, Charles suffered a crippling stroke which culminated in his early retirement. This unfortunate event disrupted his plans for completing the key project, although he has continued to work as a Research Associate of Landcare Research. In May 1994 Jan Klimaszewski arrived from Canada to work as a taxonomist on the beetle fauna of New Zealand. His primary task was to check and update Charles Watt's key, but he opted to prepare a manuscript reviewing New Zealand families, using many of Charles Watt's examples of species. In consultation with Charles this was completed in late 1996, when the manuscript was accepted for inclusion in the n f l nd series. As first author, Jan was responsible for preparing the manuscript, the present concept of the families, the family descriptions and their arrangement according to the most recent system of classification, the data on nomenclature, geographic distribution, and economic importance, the selected references, and line illustrations of structures. He also produced a second, pyramidal key intended for quick identification. In late 1996 Jan took up an opportunity for employment in Canada, so as to be reunited with his wife Krystyna and son Philip. The published version has been considerably improved by the suggestions of several reviewers and collaborators, all of whose input is warmly acknowledged, and with significant support and guidance from the n Management Committee (Marie-Claude Larivière, Trevor Crosby, Ross Beever) and the Invertebrate Systematics Advisory Group (see p. ii). It has been structurally developed and completed by the Series Editor, working from Jan's final draft, and integrating the many suggestions of reviewers and collaborators. This contribution attempts to fill a long-felt need, and critical feedback from users is invited, in anticipation that a revised edition may incorporate many further improvements.
—Tymone Duval, Series Editor, June 1997 WI OU A O OMY O GI E S A E O E ICKS A SYS EMA ICS O E US OW O U EM OGE E E OUSE O IO OGICA SCIE CE IS A MEA I G ESS UM E
— R.M. May, 1990: Taxonomy or destiny. t r 4 : Ι 9-13
E ICA IO
h r r p tf ll d d t d t G K h l (1 r n ( nd h 2)3 A t n r ( r nd-br n r n b tl f l nd nd A tr l nd n b tl l f t n n n r l n r d th ntr b t n nd n bl d t r l t n
(1 r G ll r (`W ll K h I n r t r d r r h nt t n C I pt r n th f r r Ent l v n f S1 n t ll t l n nd l t r t Mt Alb rt A I nd l r I r p n bl f r th xp n n f th SI n t ll t n r l nt f th l nd Arthr p d C ll t n W ll d I n n nd r p t d f r h r r h n th t t f C r l n d
( r hn r n Ch f r h S nt t th CSI O nd C r t r f C l pt r t th A tr l n t n l In t C ll t n C nb rr hn n t t nd n l pt r t nd rld l d r n th h h r l f t n f b tI r n A tr l n b tl nd h n t f l f t n n bl d t pl t r pr nt ntr b t n
(3 r AIfr d t n r n A t C r t r nd d f th v n f In t t th ld M Ch U S A nd r n n d p l t n th h h r l f t n f St ph l n d Al nt r t d n th rld f n f b tl nd n n p rt nt p p r h p bl h d n th l nd nd A tr l n f n
— Jan Klimaszewski & Charles Watt A S AC The 82 beetle families represented in New Zealand are reviewed. These families are briefly characterised in the phylogenetic order proposed by Lawrence & Newton (1995),1 with family
concepts .similar to those defined by these authors and by Pakaluk t l (1994). 2 A general introduction to the New Zealand beetle fauna is given, and collecting methods are briefly discussed, and core literature is reviewed. Two alternative keys—pyramidal and linear—are provided for identification of the families, using often different character sets. Morphological features are explained, and many are illustrated. Estimated numbers of species, genera, and (where appropriate) tribes and subfamilies are given. Known distributions are discussed, and data on natural history and collecting are summarised. For each family at least one species is represented by a habitus illustration, short description, and geographic profile, and important references are listed. Keys are provided to subfamilies of Carabidae, Leiodidae, Staphylinidae, Trogossitidae, Phloeostichidae, Scarabaeidae, Cerambycidae, Chrysomelidae, Anthribidae, Belidae, Brentidae, and Curculionidae; for some other families a substantive publication containing a key is cited. Some 1100 currently recognised genera are recorded for New Zealand, and approximately 5580 species (>5223 native, >3546 adventive) are recognised, including undescribed species housed in the New Zealand Arthropod Collection. One endemic family, Chalcodryidae, and four endemic subfamilies—Horelophinae (Hydrophilidae), Euderinae (Bostrichidae), Agapythinae (Phloeo- stichidae), and Cyclaxirinae (Phalacridae)—are reported. An undescribed species f l t from the Three Kings Islands is the only representative of the family Cucujidae in New Zealand. 1 l nd bf l f C l pt r ( th l t d n r n t r f r n nd d t n f l - r p n In: l A Sl p n ( d l ph l n nd l f t n f C l pt r p p r l br t n th th b rthd f Cr n W r z M In t t t l A p 779-1 2 C rr nt l f t n nd f l - r p n n C j d (C l pt r Genus 5(4): 3-
C ECK IS O AMI IES The checklist sequence follows mainly the classification Superfamily STAPHYLINOIDEA proposed by Lawrence & Newton (1995). Numbers in [7] Family Hydraenidae 5 32 0 24 brackets are unique identifiers for each family. Columns of [8] Family Ptiliidae 13 >48 8/9 24 numbers are estimated totals of genera and species [9] Family Agyrtidae 1 2 0 2 currently recognised in New Zealand: col. 1 – genera; col. [ 0] Family Leiodidae 2 112 1 2 2– native species; col. 3 – adventive species. Numbers of [11] Family Scydmaenidae 11 >201 1 26 species are based on published data (e.g., Holloway 1982, [12] Family Staphylinidae 0 >936 >85 26 Watt l982a, 1992, Ordish 1984, Werner & Chandler (incl. Microsilphinae, Pselaphinae, Scaphidiinae) 1995, Klimaszewski t l 1996) and the total number, including undescribed species, in the New Zealand Arth- Series SCARABAEIFORMIA ropod Collection (NZAC). Superfamily SCARABAEOIDEA [13] Family Lucanidae 7 26 3 2 Suborder A E AGA Page [ 14] Family Trogidae 1 0 1 0 Superfamily CARABOIDEA [15] Family Scarabaeidae 2 132 12 0 [ ] Family Rhysodidae 4 6 0 20 [2] Family Carabidae 75 >426 >19 2 (incl. Cicindelinae) Series ELATERIFORMIA [3] Family Dytiscidae 11 16 0 22 Superfamily SCIRTOIDEA [4] Family Gyrinidae 1 0 1 22 [16] Family Scirtidae 11 >125 0 31 (= Helodidae) Suborder O Y AGA [17] Family Eucinetidae 1 1 0 2 Series STAPHYLINIFORMIA [18] Family Clambidae 2 9 2 2 Superfamily HYDROPHILOIDEA [5] Family Hydrophilidae 2 70 5 2 Superfamily BUPRESTOIDEA [6] Family Histeridae 13 23 6 2 [ 19] Family Buprestidae 2 2 1 33 -9- S p rf l Y OI EA [57] l C r l ph d 8 19 2 4 [ ] l Byrrhidae 8 79 0 33 [5 ] Family Corticariidae 1 53 11 55 [ 1] l Dryopidae 2 4 0 4 (f r rl as Lathridiidae) [ ] l El d 1 1 0 4 [ 3] l Limnichidae 2 7 1 35 S p rf l E E IO OI EA [ ] l Heteroceridae 1 1 0 35 [59] l Mycetophagidae 3 1 3 6 [ 5] l Ptilodactylidae 2 - 0 35 [ ] l Archeocrypticidae 1 0 1 6 [ ] l Chelonariidae 1 1 0 3 [ 1] l C d 5 20 0 6 [ ] l Melandryidae 9 3 0 57 S p rf l E A E OI EA [ 3] l Mordellidae 4 6 1 57 [ 7] l Eucnemidae 7 22 0 6 [ ] l Rhipiphoridae 3 5 0 8 [ ] l Elateridae 26 13 3 37 [ 5] l C l d d 26 19 0 8 [ 9] l d 1 0 1 37 [ ] l Ulodidae 4 20 0 59 [3 ] l Cantharidae 3 40 1 8 (f r rl ph r d [ 7] l Chalcodryidae 3 5 0 59 S r OS IC I O MIA [ ] l Tenebrionidae 3 139 1 60 S p rf l E O O OI EA [ 9] l Prostomidae 1 1 0 6 [31] Family Derodontidae 1 2 0 39 [7 ] l Oedemeridae 6 1 3 6 [71] l Pyrochroidae 3 >7 0 62 S p rf l OS IC OI EA [7 ] l Salpingidae 4 22 0 62 [3 ] l Jacobsoniidae 2 3/ 0 39 ( n l In p pl n [33] l Nosodendridae 1 /3 0 40 [73] l Anthicidae 7 17 9 6 [3 ] l Dermestidae 7 11 6 40 [7 ] l Aderidae 2 15 0 64 (f r rl E l n d [35] l Bostrichidae 4 1 7 4 [75] l Scraptiidae 2 >4 0 64 ( n l t n [3 ] l Anobiidae 5 28 11 42 S p rf l Cn YSOME OI EA ( n l t n n [7 ] l Cerambycidae 6 1 8 6 [77] l Chrysomelidae 13 19 66 S r CUCU I O MIA ( n l r h n S p rf l C E OI EA [37] l Trogossitidae 10 >24 1 4 S p rf l CU CU IO OI EA [3 ] l Chaetosomatidae 2 4 0 44 [7 ] l Nemonychidae 1 4 0 6 [39] l Cleridae 8 37 2 44 [79] l Anthribidae 28 5 3 68 [ ] l Phycosecidae 1 1 0 4 [ ] l Belidae 3 11 0 6 [ 1] l Melyridae 4 33 0 4 ( n l A l d r n = r t rh n n ( l t d [ 1] l Brentidae 3 3 1 6 ( n l Ap n n S p rf l CUCU OI EA [ ] l Curculionidae 2 1 9 46 0 [ ] l Nitidulidae 1 1 11 46 ( n l S l t n [ 3] l Monotomidae 2 1 5 46 Totals ( ppr x 1100 > 22 > 6 ( n l h z ph n [ ] l Phloeostichidae 2 2 0 4 [ 5] l Silvanidae 8 7 5 4 CO E S [ ] l Cucujidae 1 1 0 48 A n l d nt 11 [ 7] l Laemophloeidae 2 3 3 4 Intr d t n 1 [ ] l Phalacridae 2 3 1 4 Or n nd ff n t 1 [ 9] l Cavognathidae 2 5 0 0 C p t n f th b tl f n 13 [5 ] l Cryptophagidae 1 1 1 0 C n rv t n t t 13 [51] l Languriidae 5 8 0 51 M rph l nd b l 1 [5 ] l Erotylidae 2 9 0 51 St d nd d nt f t n f b tl 17 [53] l Bothrideridae 2 6 1 2 E p nt nd ll t n th d 17 [5 ] l Cerylonidae 2 5 1 2 A n th l t r t r h lpf l r f r n 1 [55] l Endomychidae 3 6 1 53 How t use this r v 19 ( n l M r ph n v f f l 20 [5 ] l C n ll d 19 22 1 4 f r n 7
— 0— Glossary of technical terms 91 lionoidea and subfamilies of Curculionidae, and a list of Keys to taxa: I `Pyramidal' key 97 genera for Curculionidae and Cerambycidae, and assisted II `Linear' key 105 in gathering data on many families. Beverley Holloway Illustrations: Fig. 1-100, schematic figures 113 contributed to sections on Lucanidae and Anthribidae. Fig. 101-260, habitus figures 128 Rowan Emberson and John Marris (Lincoln University, Examples of species (expanded captions) 168 Canterbury) contributed their knowledge to improve Appendix 1: Threatened species of N.Z. beetles 184 various parts of the text, and provided many distribution 2: Beetle collections in N.Z. 184 records for the families. Rowan also assisted in correcting 3: Key to tribes of Carabidae 185 the difficult sections on Carabidae and Scarabaeidae. John 4: Distribution records for families 186 Lawrence (CSIRO, Canberra, Australia), Al Newton, Jr Taxonomic index 188 (Field Museum, Chicago, U.S.), and Margaret Thayer were extremely helpful in providing unpublished data, sharing their knowledge, and commenting on the manu- ACK OW E GME S script. Laurent LeSage (Agriculture Canada, Ottawa, Canada), Richard Leschen (Michigan State University, This work is a compilation from previous taxonomic East Lansing, U.S.), Jyrki Muona (Finnish Museum of revisions, occasional short papers, and beetle specimens Natural History, Helsinki, Finland), Masahiro Ohara that have accumulated in the New Zealand Arthropod (Hokkaido University, Japan), Stewart Peck (Carleton Collection over an extensive period. The specimens were University, Ottawa, Canada), and Clarke Scholtz (Uni- collected primarily by G. Kuschel, J.C. Watt, J.S. Dug- versity of Pretoria, South Africa) provided help and dale, B.A. Holloway, B.M. May, and many others not valuable suggestions. We express our appreciation to mentioned but whose efforts deserve our sincere thanks. these collaborators, and to the many others who directly or Des Helmore executed the great majority of the habitus indirectly contributed to our work. drawings. His superb work is indispensable to us and to the For critical comments and helpful suggestions on users of this publication; we salute him for his contri- various parts of the manuscript we thank Barbara Barratt bution. The work of other contributing illustrators— (AgResearch, Invermay), John Bradshaw (Department of Anthony Harris, Joanna Liddiard, Shaun Forgie, and Geology, University of Canterbury), George Gibbs Lesley Alexander—is warmly acknowledged. We also (Victoria University, Wellington), Anthony Harris (Otago appreciate the permission of John Lawrence and Jarmila Museum, Dunedin), John Hutcheson (Forest Research Kukalová-Peck to reproduce figures first published in Institute, Rotorua), Chris Reid (James Cook University, Lawrence & Britton (1994) and Kukalová-Peck & Queensland, Australia), and Greg Sherley (Department of Lawrence (1993). Formal acknowledgment for illus- Conservation, Wellington). trations is made in the captions (see especially p. 127). We are extremely grateful to the editor of the Fauna Colleagues at Mt Albert Research Centre read the first series, Tymone Duval, for his enormous assistance in draft and improved it with their comments: our thanks editing, formatting, and improving the final version of the to Ross Beever, John Dugdale, Dianne Gleeson, and manuscript. His contribution was essential to achieve the Research Associates Willy Kuschel and Brenda May. We present level of data organisation. also thank Ross Beever for promoting an atmosphere of Jan Klimaszewski would like to thank his wife Krystyna trust, creativity, optimism, and quiet scholarship. Marie- for her moral support and understanding during the pre- Claude Larivière assisted with the computer literature paration of this work. search on staphylinids, and provided the initiative and Charles Watt takes this opportunity to renew his technical savoir faireto permit in-house scanning of the gratitude to Dr Roy A. Crowson (University of Glasgow) habitus illustrations. Trevor Crosby helped in many ways for hospitality and intellectual stimulation. Many years during the preparation of the manuscript; his influence has ago Dr Crowson visited New Zealand, and inspired greatly improved the introductory sections and the Charles's interest in the family classification of our references. Birgit Rhode checked and corrected German beetles. When Charles was studying at Oxford on the references and their translations. André Larochelle helped relationships of tenebrionid beetles Dr Crowson invited in organising major publications on our beetles according him to Glasgow, where he was able to study extensive to family; he also assisted in gathering data on his collections of New Zealand tenebrionid larvae, and specialty, the Carabidae, and checked and corrected the benefited from evening discussions on beetle family French references and their translations. The friendly staff classification. Charles's manuscript linear key to the of our Centre Library—Marleene Boyd, Jillian Irwin, and families of New Zealand beetles—circulated before he Bella Smith—provided assistance with the references. went on study leave to Europe in 1984—was based on We are most obliged to Willy Kuschel, who shared with Crowson's classification as it existed at that time. us his extensive knowledge of New Zealand beetles, Charles wishes to especially acknowledge his father, the provided keys to identification for families of Curcu- late Dr John S. Watt, who on learning of Charles's stroke in 1985 flew to Paris and remained with him until he was instance the relict genera of tenebrionoid Pilipalpinae per- able to be flown home. When Charles had recovered sisting in New Zealand, Australia, southern South Amer- sufficiently, his father began to bring him twice weekly to ica, and Madagascar (Pollock 1995). The Chaetosomatidae the Mt Albert Research Centre, where they would sit are known only from New Zealand and Madagascar, and together preparing some of Charles's earlier work for the Phycosecidae from New Zealand, Australia, New publication. This continued until John Watt, well into his Caledonia, and Vanuatu (J.F. Lawrence, in litt.). The small eighties, was himself laid low by a stroke, and died in late family Cavognathidae, with species occurring in birds' 1996. The role of helper has been taken up by Charles's nests, is known from Australia, New Zealand, and Chile. sister Mrs Heather Raudon. Charles is profoundly grateful The New Zealand Nemonychidae are closest to species for this help, which was essential for him to be able to from Chile. Omaliine staphylinids of the genus Metacor- complete his part of the present work from a wheelchair neolabium (22 species) are distributed in New Zealand, and with impaired visual acuity. Australia, and South America (Thayer 1985). Some south- This work was supported by the Foundation for ern temperate Staphylinoidea from families Hydraenidae, Research, Science and Technology under contract nos Ptiliidae, Agyrtidae, Leiodidae, and Staphylinidae (esp- C09308 and C09617. ecially Omaliinae and Pselaphinae) show `transaustral disjunctions,' i.e., are isolated in two or more southern land areas—New Zealand, Australia, South Africa, and/or South America; for details see Newton (1985). In contrast, I O UC IO the small, archaic family Agyrtidae, with fossil records from the mid Jurassic (Newton 1991a), is distributed in the Beetles are the largest order of organisms, comprising over Holarctic region except for two species occurring in New 350 000 described species in the world fauna. They are Zealand. Similarly, the staphylinid Stylogymnusa subant- classified in some 23 000 genera (Endrödy-Younga 1985). arctica is known only from the Auckland Islands, with its Lawrence & Britton (1994) estimated that there are more closest relatives exclusively Holarctic. species of beetles than there are vascular plants or fungi, The factors responsible for the composition and and 90 times as many as mammal species. For a com- diversity of the present-day New Zealand beetle fauna are: prehensive definition of beetles see Lawrence & Britton (1) the Gondwana origin of some of our biota; (2) the (1994). estimated 80 million years of geographic isolation of New The first collections of New Zealand beetles were made Zealand, which resulted in some 90% endemism at the by Joseph Banks and Daniel Solander during Cook's first specific level and a high level of generic endemism; (3) voyage in 1769, and the first species were described by changing climate, changing shorelines (Oligocene bottle- Fabricius (1775). Watt (1979b, 1982a) and Ramsay & neck), progenies, glaciation, and volcanic activity; and (4) Singh (1982), amongst others, provide biographical notes the absence of mammals and other animals and plants about the early collectors and describers of beetles, and the which have dominated ecosystems elsewhere in the world. development of entomology in New Zealand. The fauna has been greatly influenced in the last 1000 years by humans, who brought devastation to native forest Origins and affinities and introduced exotic animals with destructive influence The exact origin of New Zealand beetles is essentially (e.g., deer, rats, possum, goat). unknown, but there are indications that at least some Contemporary New Zealand beetles are a composite of beetles had developed on an ancient landmass of con- ancient, variously changed lineages, elements introduced tinental origin which became increasingly isolated since by dispersal over short and long distances (e.g., groups the late Mesozoic (Daugherty et al. 1993). For example, dispersed passively over water as a result of the strong the New Zealand jacobsoniid beetle Saphophagus min- westerly winds and currents around the 40th parallel), and utus is considered by Crowson (1959) to be a relict coeval species intentionally and accidentally introduced by with the tuatara (Sphenodon). humans. In support of the hypothesis that at least some of Craw & Watt (1987) reported the first New Zealand our beetles are of ancient origin is the extraordinarily high Cretaceous beetle fossil, an incomplete elytron, identified level of endemism at the specific level, with several as belonging to a member of Polyphaga, either a buprestid endemic tribes, some endemic subfamilies, and many or a chrysomelid. Kuschel (1987b) described subfossil endemic genera (Watt 1982a). Several families usually fragments 1680 years old of a presumably extinct fern- small elsewhere are represented in New Zealand by weevil, Tymbopiptus valeas, from near Te Kuiti, North diverse endemic forms (e.g., Byrrhidae, Colydiidae, Island. For a recent review of the geological context of the Hydraenidae, Melandryidae, Ptiliidae, Scirtidae, and evolution of the New Zealand biota, see Cooper & Scydmaenidae). The native element in our fauna is Millener (1993). associated primarily with the lowland forest which has In some groups present-day distribution patterns are prevailed throughout most of New Zealand's geological suggestive of a Gondwana or pre-Gondwana origin, for history (Kuschel 1990), alpine habitats, tussock grass-
-12- lands, and the subantarctic islands. Introduced species are The most comprehensive survey ever conducted on mostly associated with man-induced habitats (Kuschel New Zealand beetles is that of Kuschel (1990), in the 1990). suburb of Lynfield, Auckland, in which 982 species are listed (the total number from the survey area, including Composition of the New Zealand beetle fauna some collected after the survey was published, reaches 1000 — G. Kuschel, pers. comm.). Of these, 753 are en- At least 5223 native and 356 adventive species of beetles demic and 229 adventive, and they belong to 65 families. have been recognised in New Zealand, belonging to some Of the 753 indigenous species, approximately 10 are 1100 genera in 82 families (estimates based on currently restricted to beaches or intertidal wrack and 2 others to recognised species housed in the New Zealand Arthropod beaches and open field situations. The great majority were Collection and various museums: Watt 1976, 1982a, associated with native bush. Sixty-eight species, con- Klimaszewski et al.1996). The actual number of New stituting 9% of the 744 native non-littoral species in the Zealand beetle species, including undescribed ones, is survey, were found in open environments but their sub- probably much higher. By comparison the vascular plant sistence in these environments and their endemic status flora includes about 2500 species, and the terrestrial were uncertain. Kuschel (1990) presented interesting vertebrate fauna about 350 species (Watt 1976). Watt observations on the ecological flexibility of indigenous (1976) estimated at 20 000 the number of described and (native) and foreign (introduced/ adventive) species. Of undescribed species of terrestrial (including freshwater) the 982 Lynfield beetles, 55 belong to 16 genera with both Arthropoda probably occurring in New Zealand. The native and foreign species. Of these 55 species, 22 natives numbers are estimates, but nevertheless show that beetles were found only in the native bush, and all 33 foreign constitute a significant proportion of New Zealand's total species occurred in home gardens and fields, but 14 of insect fauna. these occurred also in the native bush environment. The New Zealand beetles belong to two suborders, Kuschel (1990) believes that the beetle population of Adephaga and Polyphaga. The suborders Archostemata Lynfield represents about 8-8.5% of the total (described and Myxophaga, although present in Australia, do not and undescribed) New Zealand beetles—some 10 000 to have representatives in our fauna. Of the 82 New Zealand 10 500 species. The conclusions of the Lynfield study families, four—Archeocrypticidae, Gyrinidae, Lycidae, relate to all our beetles, which are predominantly forest- and Trogidae—have only adventive species present. inhabiting species and show little interest in man-induced Eleven families are represented by more than 100 des- environments. cribed species. They are here listed in descending order, Ground-breaking investigations on the composition of with approximate numbers in parentheses: Curculionidae beetle faunas in native forests and in degraded habitats, (1542), Staphylinidae including former Pselaphidae and of beetles as indicators of habitat degradation, are (1021), Carabidae (445), Scydmaenidae (202), Colydiidae being conducted by entomologists of the Forest Research (196), Cerambycidae (188), Chrysomelidae (153), Scar- Institute in Rotorua (Hutcheson & Hosking 1994; Hutch- abaeidae (144), Elateridae (135), Tenebrionidae (159), eson 1990, 1996; Hutcheson & Kimberley, submitted). and Scirtidae (125). The numbers of Curculionidae, Staphylinidae, and Carabidae in relation to the total num- ber of beetle species in New Zealand are similar to the Conservation status proportions in Australia and in the temperate Holarctic. The majority of New Zealand's beetle species are as- However, the Colydiidae in New Zealand are particularly sociated with native forest, a habitat which has been diversified, being better represented than elsewhere. Some drastically reduced by human activities over the past smaller families, e.g., Byrrhidae, Cleridae, Hydraenidae, millennium. Native forest now constitutes only 23% of the Corticariidae (=Lathridiidae), Leiodidae, Melandryidae, total land area. The `shrinking' forest has profoundly and Trogossitidae, are also well represented in New Zea- reduced the population size of many native species and land. There is one endemic New Zealand family, Chalco- their potential for long-term survival. Almost certainly dryidae, and a few endemic subfamilies—Horelophinae many species have vanished before being documented, (Hydrophilidae), Euderinae (Bostrichidae), Agapythinae and this process is probably continuing. Our beetles are (Phloeostichidae), and Cyclaxyrinae (Phalacridae). In also vulnerable to introduced organisms because they have comparison, there are approximately 20 000 described evolved in isolation, and their defence mechanisms are beetle species in Australia (total beetle fauna estimated at often ineffective against the new arrivals (e.g., comp- 30 000) in 117 families; the largest are Curculionidae etition for food with introduced wasps, predation by (6000), Scarabaeidae (3000), Chrysomelidae (3000), mammals, depletion of food sources by weed competition Carabidae (2500), Staphylinidae including Pselaphinae with native plants). (2500), Cerambycidae (1500), and Tenebrionidae (1200) The major threats to biodiversity are fragmentation and (data from Lawrence & Britton 1994 and Klimaszewski et degradation of ecosystems due to inappropriate use of land al. 1996). (e.g., deforestation) and water, invasion by exotic plants
—13— and animals (Ramsay 1978), non-sustainable use of The capsule dorsally consists of vertex, frons, and cly- species, and climatic changes (global warming, etc.). Con- peus (Fig. 1, 6); anterior and posterior tentorial pits, the sequently populations associated with small, fragmented former (often called just tentorial pits) located at the and degraded communities are particularly vulnerable, border of the clypeus and the frons; and ventrally the gula, and are in need of conservation. The following citation genae, submentum, and mentum. The head is usually pro- from Kuschel (1990) reflects the most important gnathous, i.e., with mouthparts facing forward (Fig. 1, 8), conclusion from his study of the beetle fauna of the but may be hypognathous with mouthparts facing down- Auckland suburb of Lynfield. ward (Fig. 9, 10). In many Curculionoidea the frons and vertex are prolonged anteriorly to form a rostrum (Fig. 6, f M h th n t v f r t f n ll r pt 7). l v nd n p r r r f r t rt n Compound eyes are usually well developed, except in l nd b f r th rr v l f th l n n some species living in permanent darkness (e.g., in deep nd l t r th E r p n h r n n th soil or in caves). They may be coarsely or finely faceted. t r xt n t n nl th nd n v t t n They are variable in size and shape (Fig. 8-12), and h h nt n th p l h b t t th t r v t l sometimes (e.g., Gyrinidae, Tenebrionidae) each eye is t th r rv v l pr rv d A th rv d n- partially or completely divided into two by a lobe (Fig. 11) tr t d v n th r l t v l ll b h p t h th t called a canthus (Nichols 1989), ocular canthus, or genal f r h v p d d tr t n nt n f n f canthus (Watt 1988). n p t d r hn nd d v r t The mouthparts (Fig. 4) are a labrum, two mandibles, Examples of small and isolated beetle populations are two maxillae each comprising a lacinia, a galea, maxillary an undescribed l t species known only from Great palps, glossae, and paraglossae, and a labium with labial Island in the Three Kings group, G d r th n palps (Fig. 5). The glossae and paraglossae are usually from Stack H in the Mokohinau Islands, and St l - fused to form a median ligula. Variations in the terminal n b nt r t occurring exclusively on the Auck- segment of the maxillary palp (Fig. 13-16) have dia- land Islands. The first is the only representative of the gnostic value for some groups. family Cucujidae in New Zealand, and the last is the only The antennae are usually 11-segmented, with the basal surviving gymnusine staphylinid (see p. 12) in the segment, or scape, articulated to the head and the second Southern Hemisphere. segment, or pedicel, connecting the scape with the rest of The Department of Conservation has provided a list of the antenna, called the flagellum (Fig. 1, 6). The last few threatened species for New Zealand (Molloy t l 1994) — segments may be swollen, forming a more or less compact see Appendix 1. There are 26 native invertebrates, in- club. The main antennal types are: cluding beetles, in category A (highest priority threatened f l f r — the simplest form, with cylindrical segments species), 52 in category B, and 20 in group C. The total which are nearly uniform in shape (Fig. 17, 18); number of threatened invertebrates represent less than n l f r — with round, similar segments, resembling a 0.1% of the taxonomic groups in categories A—C (Molloy string of beads (Fig. 19); t l 1994). Examples of New Zealand beetles of in- rr t — with one side of each segment slightly pro- determinate conservation status that are rare in collections truding, and forming a series of points like the teeth of are r n th r (Chelonariidae), n x r nd a saw (Fig. 20, 21); (Monotomidae), and species of M r lph (Staphylinidae) p t n t — with one side of each segment protruding, and r l ph (Hydrophilidae). forming a series of long, sharp points like the teeth of a comb (Fig. 22) (if both sides of each segment are prolonged, the antenna is called b p t n t pl — similar to the bipectinate antenna, but with the MO O OGY A IO OGY projections on both sides of each segment slender and flexible, like the vanes of a feather; The basic morphological structures of beetles referred to l v t — with apical segment's enlarged into a club-like in this contribution are shown in Fig. 1-3, which are expansion, either weak and gradual or abrupt and ball- intended to be self-explanatory. We will, however, briefly like (Fig. 23-29); characterise the major morphological features of the beetle p t t (a type of clavate antenna) — with apical segments body. Readers are also recommended to consult the abruptly enlarged into a ball-like club (Fig. 23, 187); Glossary (p. 91) for definitions of morphological terms. l ll t — with club segments having flat, leaf-like projections on one side which usually can (Fig. 31) or d (Fig. 4-33). The head consists of a sclerotised cap- sometimes cannot (Fig. 30) be brought closely to- sule, mouthparts, two compound eyes and sometimes one gether; or two ocelli (e.g., Staphylinidae - Omaliinae, Derodon- fl b ll t — with long, thin lateral processes like alternate tidae), and a pair of antennae. folds of a fan (Fig, 32, 33);
4 n l t – with the second segment (pedicel) attached at W n (Fig. 46-51). The first pair of wings are sclerotised an angle to the first (scape), and each following seg- to form two l tr which usually completely cover the ment in line with the pedicel (Fig. 28, 140-143). abdomen (Fig. 1) or which may be reduced, exposing most of the abdomen, as in the majority of Staphylinidae (e.g., h r x (Fig. 1-3, 34-45, 52-56). The thorax consists of Fig. 132-138). The second pair of wings are the mem- three parts—prothorax, mesothorax, and metathorax— branous flying wings (Fig. 46, 47), which are sometimes each bearing a pair of legs lateroventrally. The mesothorax small or even absent, and are concealed by the elytra when and metathorax are joined together to form the ptero- not in use. For details on venation and evolution of the thorax, and each bears a pair of wings dorsolaterally. hind wings in Coleoptera (Fig. 48-51) see Kukalová-Peck Each thoracic segment is made of four sclerites, with & Lawrence (1993). their borders defined by sutures. The topmost (dorsal) sclerite is the notum, the two lateral sclerites are the pleura, (Fig. 1, 57-87). The beetle leg consists of a x and the bottom one (ventral) is the sternum. Any one of the (usually with tr h nt n trochanter, a f r a t b thoracic sclerites is named by adding the appropriate and a t r The coxa of each leg articulates with the prefix: hence pronotum, propleuron/-pleura, prosternum, thorax at two points: the trochantin, which is the only mesonotum, mesopleuron/-pleura, mesosternum, etc. The externally visible extension of the internal pleuron (Fig. prominent part of the beetle thorax is the pronotum, which 57), and the pleuron, which is separated from the troch- is variably shaped and is often referred to as the pronotal antin by a membrane. The trochanter is articulated to the disc (Fig. 1). coxa (Fig. 58-66). Its junction with the femur may be In Adephaga (also Archostemata and Myxophaga, not straight (Fig. 61) or slightly oblique (Fig. 60), but some- represented in New Zealand) the pronotum is extended times is strongly oblique, as in Fig. 58 and 59 (e.g., in most and deflexed ventrally on either side, forming the pr n t l Tenebrionoidea, Bothrideridae), so that there is direct p pl r n (Fig. 2, 34, 35). The pronotal epipleuron meets contact between the femur and coxa (h t r r d the lateral part of the pl r n (pr pl r n which is trochanter). Each tarsus is divided into 1-5 segments visible externally and forms part of the thoracic wall. The (tarsomeres or tarsites), and the last segment usually bears pleuron (propleuron) is separated from the pr n t l p - two claws (Fig. 67-82), rarely one. The tarsal formula pl r n by the n t pl r l t r and from the t rn refers to the number of tarsal segments on each pair of legs. by the pl r t rn l t r It may be divided into two sclerites, an anterior pr p t rn which forms part of Abd n (Fig. 2, 3, 88-92). The abdomen consists the wall, and a posterior pr p r n which may be apparently of 10 segments in the male and 9 in the female, extended to meet the sternum and closes (partially or each forming a complete or partial ring. The 9th segment is completely) the coxal cavities from behind. The tr h- modified, forming the genital segment, and in the male the nt n is a small sclerite separated (or not) from the pleuron 10th segment is usually much reduced or fused to segment by a membrane and connected by the articulating condyle 9. Each abdominal segment is composed of four sclerites: with the coxa (Fig. 57). a dorsal tergum or tergite, two pleura or pleurites (one on In Polyphaga the deflexed lateral portion of the pro- either side), and a ventral sternum or sternite. A respiratory notum is called the h p r n and is attached directly to spiracle opening onto each pleuron is the aperture of a the lateral portion of the sternum because the pleuron has trachea. The spiracles are usually located in pleural mem- become reduced, fused with the trochantin, and hidden brane, but in some Scarabaeoidea and Staphylinoidea they internally ( nd pl r n (Fig. 3, 36, 37). The hypomeron may be located on tergites. is separated from the sternum by the n t t rn l t r The visible part of the abdomen consists of five or more the n t pl r l t r is absent as a consequence of sclerotised t rn t (called v ntr t (Fig. 2, 3). The first reduction and internalisation of the pleuron. The anterior sternite is absent, or represented by a small sclerite near the portion of the sternum varies among Polyphaga. metacoxae; the second sclerite may be hidden behind the The nota of the mesothorax and the metathorax are edge of the elytra, and is visible only in lateral view. In divided into three sclerites: the t t ll and referring to the first sternite we mean the first ventrite or p tn t The mesoscutellum, usually called just the the first visible sternite. In Adephaga the first ventrite is scutellum, is visible at the base of the elytra (Fig. 1). The completely divided by the large and prominent metacoxae ventral part of the pterothorax consists of two sterna and (Fig. 88, 89), but in the Polyphaga it is undivided and the pleura ( p t rn and p r (Fig. 2, 3). The coxal nt n fr d t d ( ί 9 h f r t v ntr t cavities are bounded anteriorly and posteriorly by the may bear a pair of straight or rv d f r l l n behind sternum and laterally by the pl r l l r t The meso- or between the metacoxae (e.g., in Coccinellidae) (Fig. 92, coxal cavities may be considered as laterally `open' when a). The abdominal ventrites are either movable and partly enclosed by pleural elements (Fig. 42, 44) and separated from each other by an intersegmental membrane laterally `closed' when closed by the sterna alone (Fig. 43, (Fig. 91), or some of them are nn t lacking an 45). intersegmental membrane but with the suture still visible
– 15 – Table 1 Primary feeding mode of some beetle families Pupae are partly enclosed by the larval skin in surface- and subfamilies (adults and larvae). feeding beetles, or pupation occurs within the food plant or in cells in the soil. Some species make cocoons, which in Predators certain exotic aleocharine staphylinids are constructed Cantharidae, Carabidae, Cleridae, Coccinellidae, Coly- from silk (Ashe 1982). diidae, Cucujidae, Lycidae, Staphylinidae, Trogossitidae Coleoptera larvae have a well developed and usually Phytophages sclerotised head, three thoracic segments, and commonly Apioninae, Chrysomelidae, Curculionidae, Nemonychidae, ten abdominal segments (rarely eight or nine). The first Nitidulidae, Scarabaeidae thoracic segment is slightly enlarged and more heavily sclerotised, abdominal segment 9 usually has paired Scavengers r fungivores processes called r ph and the 10th abdominal Anobiidae, Anthribidae, Chalcodryidae, Ciidae, Corticari- segment is small and sometimes bears p p d Beetle idae, Cryptophagidae, Dermestidae, Lucanidae, Ptiliidae, larvae usually have mouthparts adapted for biting and Ptininae, Tenebrionidae chewing, as in the adults, and have usually from three to six simple eyes, called ll or t t (Lawrence 1991). The larval body varies a lot but is usually cylin- (e.g., Buprestidae). Occasionally the suture between the drical, and is either legless or with apparent appendages. first two visible segments may be partially obliterated. The Many larvae which bore into plant tissue lack legs (e.g., in last tergite of the abdomen when exposed forms the Curculionidae). Although beetle larvae are morphologically p d The reproductive organs are usually concealed very diverse, their mode of life has led to the development within the tip of abdomen. of a few distinctive larval types; for details and illus- The internal genital structures, although very important trations see Lawrence & Britton (1994). for distinguishing species, are complex and are not dis- The main morphological categories of larvae are cussed here; readers may consult Lawrence & Britton r f r r b f r p d and p d f r (1994). (Fig. 97-100). Eruciform larvae are cylindrical or some- what flattened, with short legs and with either short uro- gomphi or none. They are less active than campodeiform Biology larvae. Many species of Chrysomelidae and Tenebrionidae Adults usually are more conspicuous than larvae and occur have this type of larva. Scarabaeiform larvae are distinc- virtually in all terrestrial habitats; they may be nocturnal or tive by their robust, C-shaped body and Iong legs. They diurnal, and may or may not occur in the same habitat as occur in the soil or rotten wood, and are characteristic for their larvae. Kuschel (1990) discussed habitat types and Scarabaeoidea. Apodous larvae have reduced antennae the beetle fauna in the suburb of Lynfield, Auckland. The and palps and lack thoracic legs and urogomphi. This type range of habitat niches even in a suburban environment is is typical for Curculionidae. Campodeiform larvae have a overwhelming, and includes forest canopy, tree trunks, prognathous head, long and well developed legs, and com- live wood, sound dead wood, decayed wood, wood-rotting monly long, unisegmented or multisegmented urogomphi moulds, ground plants and fungi, leaf litter, compost, (Lawrence & Britton 1994). They are active and usually dung, carrion, stream banks and beds, soil, beach sand, predatory. Most Carabidae and Staphylinidae have this wrack in the intertidal zone, ponds, etc. type of larva. Beetles feed on a variety of plant and animal material, Larvae are less conspicuous than adults and occur and parts of living plants may be utilised (roots, stems, usually within their food source, e.g., inside timber leaves, and flowers). Slime moulds, fungi, algae, decaying (Cerambycidae, Buprestidae, Anobiidae), inside stems wood, leaf litter, carrion, dung, and other insects are food and leaves (Chrysomelidae, Curculionidae), in fruits sources for various beetles. Feeding modes are summar- (Nitidulidae) and seeds (Bruchinae), in the soil and/or leaf ised in Table 1. litter (Staphylinidae, Carabidae, Pselaphinae), in roots or For information about the biology of the Coleoptera in dung (Scarabaeidae), in carrion (some Staphylinidae, e.g., general, see Crowson (1981). Al h r or on the ground near the base of plants. Beetles are h l t b l insects, i.e., they undergo a Larvae of Coccinellidae occur on foliage, where they prey complete metamorphosis. Adult beetles lay eggs which on soft-bodied insects. Many beetle larvae occur under the hatch into larvae; frequently these occupy different bark of logs or in rotten wood (e.g., Silvanidae, Cucu- habitats from the adults. When the larva has completed jidae), others are associated with fungi (e.g., Ciidae, feeding and growing it enters a `resting' stage called the Cryptophagidae, Erotylidae), and some occur exclusively pupa, in which the body of the larva is transformed into in birds' nests (Cavognathidae). Many larvae are of that of an adult (metamorphosis). This enables particular economic importance because they compete for human species to use different tr ph n h (as larvae) and to food resources or damage human food products, and are prevail in unfavourable conditions (as pupae). therefore regarded as pests, e.g., larvae of some Derm-
6 estidae, Elateridae, Scarabaeidae, Chrysomelidae, Curculio- connecting it with the lower hoop. The latter is attached to nidae, Anobiidae, Cerambycidae (see Booth et al. 1990). a sleeve 1 metre long which can be tightened at the bottom. A family-level treatment f the larvae of New Zealand Sifters are useful for collecting beetles from all kinds of beetles by Dr J.F. Lawrence (CSIRO, Australia), with keys ground litter, rotting wood, fungi, shore debris, lichens, for their identification, is anticipated to be published in our mosses, leaf mould, and birds' nests. The beetles may be Fauna series. Consequently mention of larvae in this work picked from sifted material placed on white sheets or large has been kept to a minimum. white trays, or the sifted debris may be placed into a Berlese funnel or other separator.
Berlese funnels. The Berlese funnel in its various modi- S U Y A I E I ICA IO O EE ES fications is used to separate insects from sifted debris. The Important repositories of New Zealand beetle collections sifted sample with insects is placed on a screen near the top are identified in Appendix 2, with contact details. of a funnel which is closed by a cover. An incandescent bulb above the sample produces light and heat, drying the Equipment and collecting methods litter and driving the insects downwards into the funnel, which ends with a container at the bottom. The container Beetles occur in virtually all terrestrial habitats, therefore may be empty for catching live insects, or may contain a collecting them requires knowledge of many different killing agent. techniques. The most common and effective methods for collecting beetles in various environments are listed Flight interception traps. The basic principle of such below. For more details on collecting and mounting traps is to catch flying beetles by placing a screen in their specimens see Martin (1977), Steyskal et al. (1986), way. The most successful interception traps tested in New Walker & Crosby (1988), and Kuschel (1990). Zealand are the window trap, consisting of a vertical net with a trough containing ethylene glycol underneath, and Hand-collecting. Mainly medium-sized to large specimens the Malaise trap, which consists of a vertical net serving as may be collected by hand from a variety of habitats (e.g., a baffle, end nets, and a sloping canopy leading up to a from under stones, bark, rotting wood, foliage, etc.). collecting container. Attractants may be used to increase the efficiency of these traps, of which there are many Collecting nets. There are three basic types of nets: aerial varieties. nets for collecting flying beetles, sweeping nets for sweeping vegetation, and aquatic nets for gathering Pitfall traps. Pitfall traps are very successful devices for insects from water. Aerial nets are light and the most collecting ground-walking beetles, or flying beetles if a fragile. Sweeping nets are more robust, made of strong bait is used. They consist of an open-topped container material with reinforcement around the metal ring to sunk in the earth. A cover may be placed over the open top withstand being dragged through dense vegetation. Water to prevent rain flooding, but open space is required (aquatic) nets are made of metal screening or heavy scrim between the cover and the rim of the container. The trap with a canvas bag affixed to a metal rim. may be one-third filled with ethylene glycol preservative, and the bait (carrion, dung, or a specific insect attractant Beating sheets. A beating sheet is made of strong cloth e.g., molasses, beer, fruits, cheese) may be suspended attached to a frame about 1 metre square, with two pieces inside the trap above the surface of the glycol. Trapped of light wood or metal crossing each other and fitted into a insects should be collected regularly from the glycol pocket at each corner of the cloth. The beating sheet is held solution and placed in 75% ethyl alcohol/acetic acid or secured under a shrub or tree, and the branches or mixture. foliage are beaten with a stick. Specimens will fall onto the sheet, and can be removed using fingers, forceps, or an Light traps. An effective way to collect nocturnal flying aspirator. (An inverted umbrella can also serve as a beating beetles is to use artificial light as an attractant and capture sheet.) the beetles around the light. A mercury-vapour lamp, fluorescent black light, or other lamps high in ultraviolet Sifters. Sifters are containers made of cloth, wood, or wavelengths usually give excellent results. The light plastic with a wire-mesh bottom serving as the sifter. The source may be placed against a vertically hung white sheet, size of mesh used depends on the size of the specimens with a bottom sheet spread on the ground underneath for sought. The most common model of sifter used in Europe falling beetles. More sophisticated light traps consist of a and North America is made of two hoops of metal about 30 funnel with a killing container, and a suspended light with cm in diameter, each provided with a handle. The lower vertical rigid screen most often made of transparent hoop bears a wire-mesh screen welded to its edge. The top plastic. The flying insects approach the light, hit the hoop is attached to a canvas cylinder about 30 cm long screen, and are collected in the killing container.
—17---17— Aspirator. An aspirator is a simple suction apparatus for publications form a core of early literature, and central to picking up insects. The most common model consists of a them are the seven volumes of his M n l f l nd vial of glass or transparent plastic with a close-fitting C l pt r published in four parts (Broun 1880, 1881, rubber stopper. Two rubber or plastic tubes pass through 1886, 1893b). His work is listed here n xt n even the stopper. One is attached to a rubber tube through which though it may be of only limited value to workers today. suction is created (by mouth, or with a rubber bulb) while • Chapman (1971): textbook on morphology, physiology, the other end is filtered. The second tube extends approx- and natural history of insects. imately 10 centimetres from the stopper, and sucks the insects in. This device acts as a miniature vacuum cleaner, • Cooper (1979): review of fossil Coleoptera. and is very useful for collecting small beetles. • Crosby & Larochelle (1994): preliminary list of Coleo- ptera genera and families represented in the New Zealand Fogging. Fogging with insecticides is the most recently Arthropod Collection. developed method for collecting beetles from the tree canopy, branches, trunks, or moss-covered surfaces. For • Crowson (1955): major treatise on the natural classi- details, see Erwin (1983), Allison t l (1993), and fication of the families of Coleoptera, based on detailed Lawrence & Britton (1994). morphology of adults and larvae. Purported to reflect the phylogeny of the Order, and gained wide acceptance. Killing and relaxing. Beetles should be killed in jars with Includes keys to superfamilies, families, and some sub- a little ethyl acetate absorbed into a suitable material (e.g., families. A cornerstone in the classification of beetles, coarse hardwood sawdust such as oak, or crumpled variously modified in Crowson's later papers (1960, 1967, absorbent paper) inside the killing jar. The absorbent l97la, b, 1973b). See Lawrence & Britton (1974) for material acts as a barrier between the insect and the killing literature. agent. An alternative solution is 75% alcohol with a little • Crowson (1981): textbook on biology of Coleoptera. acetic acid or commercial vinegar to prevent stiffening of • the insect's muscles. Ventral structures obscured by in- Hudson (1934): the first and only representative cata- logue of New Zealand beetles and their larvae, with many conveniently placed legs in set specimens can be exposed colour plates. Forms a watershed between the earlier by soaking the specimen for a few minutes in Barber's pioneering work and subsequent studies. An essential fluid, or water with ammonia, to soften the joints suffici- reference point. ently for appendages to be moved without damage. • Kuschel (1990): exhaustive study of the beetle fauna of A n th l t r t r : h lpf l r f r n an Auckland suburb, remarkable for identifying almost The literature on New Zealand beetles is extensive but 1000 species from native bush and modified habitats. largely fragmentary. It is therefore necessary to know how Many habitus illustrations by Des Helmore, especially of to access relevant publications. During compilation of weevils; also tabular summaries of species data, inform- references for this review it became apparent that a few of ation on collecting methods, habitat types. A primary them are of outstanding general relevance. Our selection is source of data included in the present work, and an offered here in alphabetical order, since any attempt to essential resource for the student of New Zealand's categorise in order of perceived value would be un- Coleoptera. * realistic. However, it is fair to say that a comprehensive • Lawrence (1982): synoptic review of beetle families recent bibliography, a general textbook or two on beetle with family limits and changes to classification. systematics and biology, and regional summaries both • Lawrence (1991): discussion, key, and many family general and faunistic must be almost indispensable. Any treatments of the immature stages of Coleoptera. electronic reference sources—CD-ROMs, Internet web- sites, searchable databases—must be seen as a bonus at the • Lawrence & Britton (1991, 1994): the Coleoptera chap- time of writing, though these will undoubtedly grow in ter in the primary reference book In t fA tr l and prominence. its spin-off A tr l n tl both with much relevance The following publications, then, are so relevant to the to New Zealand. present context that we have not repeated them under the • Lawrence & Newton (1982): review of the evolution family treatments. They should be borne in mind as a first and classification of beetles at the superfamily level. point of reference. Discusses origins and relationships, early fossils, sub- • Böving & Craighead (1931): the first comprehensive ordinal classification, and evolution of Adephaga and textbook on morphology of larvae. Polyphaga. • Broun (1876-1923): Thomas Broun's name is appen- • Lawrence & Newton (1995): comprehensive current ded to more species of New Zealand beetle (and, regret- classification of Recent Coleoptera at subfamily level and tably, synonyms) than any other author's by far. His above, with complete or selective list of genera included in
—18— each group. Discusses recent changes to classification, We also draw attention to the following publications with extensive reference to studies providing evidence in reviewing beetles from other zoogeographical regions. support of these changes. • AUSTRALIA: Matthews (1980-92), Lawrence & Brit- • Meads (1990): introduction to rare, endangered, and ton (1994), Semmens et al. (1992). protected invertebrates in New Zealand. • NEW GUINEA: Gressitt & Hornabrook (1977), • Neave (1939-50): the Nomenclator Zoologicus with its Gressitt & Szent-Ivany (1968). supplements is the definitive catalogue of genus-group • PACIFIC ISLANDS: van Dyke (1953), Nishida (1994), names in zoology. Κ l vá- (199 • New (1995): textbook on conservation biology of in- • NORTH AMERICA: Bousquet (1991a), Arnett (1973), vertebrates; the first to give a global overview of all major White (1983). habitats (terrestrial, freshwater, marine). • SOUTH AMERICA: Solier (1849), Sharp (1882-87), • Nichols (comp.) (1989): a revised edition of the well Blackwelder (1957). known and comprehensive Torre-Bueno glossary of ento- • SOUTHERN AFRICA: Scholtz & Holm (1985). mological terminology. • GENERAL: Evans & Bellamy (1996). • Pakaluk & Slipinski (eds) (1995): a collection of the Important references on beetle larvae are Arndt (1993), most recent papers on biology, physiology, and classi- Booth et al. (1990) (pest species), Böving & Craighead fication of Coleoptera. (1931), Hudson (1934), Lawrence (1991) (key to fam- • Ramsay & Crosby (1992): bibliography of New Zealand ilies), Lawrence & Britton (1991, 1994), Lawrence & entomology up to 1985, and a guide to the bibliographic Newton (1982, 1995), Lawrence et al. (1993) (beetle database BUGS, which is searchable on a `by family' basis larvae keyed on CD-ROM), May (1966, 1993) (larvae of [accessible on application to the Curator, NZAC, email Curculionoidae), Stehr (1991). address [email protected]]; see also Crosby & Ramsay (1992, 1994). How to use this review • Ramsay & Singh (1982): concise guide to entomology The section Review of Families is presented in systematic in New Zealand, including an historical review. sequence—notionally reflecting a natural phylogeny—of • Shapiro & Porter (1989): paper reviewing the signi- numbered families, as in the Checklist of Taxa. Groupings ficance of genitalia in the interpretation of insect evolution from suborder to superfamily are briefly introduced as a and systematics. framework to this systematic treatment. Under each family [numbered in brackets, for ease of • Snodgrass (1935): textbook on insect morphology, use- cross-reference throughout] any assistance received from ful for naming of external and internal structures. specialist collaborators is acknowledged. The next line • Walker & Crosby (1988): concise manual on the identifies the habitus figure(s) representing that family. preparation and curation of insect specimens.* Other relevant illustrations are not listed here, but appear in the text wherever appropriate. Next, the range of body • Watt (1956-92): Charles Watt's published work is listed length—measured from the apical margin of the labrum to here in full, in view of its great relevance to the modern the apex of the abdomen—for members of the family is study of Coleoptera in New Zealand. His Presidential given, along with the tarsal for-mula. This information Address to the 1981 conference of the Entomological alone can be helpful in delimiting certain families or (Watt l982a), is a Society of N.Z., New Zealand beetles groups of families. valuable review of the field and a primary source of Subsequent information for each family is listed under a information for the present work. sequence of subheads. The well defined families are ac- • Wilson (1971): review of social insects and their affili- companied by a relatively short Diagnosis and Synopsis, ations with other insects, including beetles. but those represented by more diverse forms have required • Winterbourn & Gregson (1989): guide to the aquatic longer treatments. Character states in bold type constitute insects of New Zealand, including the five or six beetle the primary diagnosis; the balance are supplementary. It families with aquatic or hydrophilous habits as adults and/ will be noted under Synopsis that the family is referred to or larvae. by the contracted form of its scientific name, as for ex- ample Carabidae / carabid(s). This informal usage may be preferred to colloquial names suggested under Remarks. Range is represented as `recorded' (signified by ®) or *AvaiIable from Manaaki Whenua Press, P.O. x n ln `unrecorded' (O) in the three major and eight minor 15 l nd t l ( 3 3 5 7 f x ( 3 3 5 1 7 l islands or island groups constituting New Zealand; it is pr l nd r r nz . shown for all families alphabetically in Appendix 4. -19- Under Examples, species representative of the family or E IEW O AMI IES constituent subfamilies are listed. Authorship and year are cited, so that original descriptions may be pursued. Figure Suborder ADEPHAGA numbers give access to both the habitus illustration and its We have followed the concept of Lawrence & Britton expanded caption (under Examples of Species), which (1994) in defining this suborder: 1st abdominal sternite offers a profile of the species illustrated. divided by metacoxae (Fig. 2); abdominal sternites 6 in Under Remarks are given any additional points of number; pygidial defence glands present; larvae char- information thought to be helpful, including reference to acteristic (for details see Lawrence & Britton 1994). The any revisions known to be in progress, followed by a great majority of species are predacious. The phylogeny suggested colloquial name. Some of these names are well and evolutionary history of the Adephaga is discussed by established internationally, others are acceptable options Beutel (1995). The Adephaga comprise four families in a in New Zealand. single superfamily, all represented in New Zealand. The major families are furnished with a Key to their subfamilies, or with references to such keys. Finally, under Selected References we offer a list of Superfamily CARABOIDEA publications considered likely to be most helpful in further This superfamily consists of families Rhysodidae, Cara- exploring the family under consideration. These have been bidae (including Cicindelinae), Dytiscidae, and Gyrinidae. kept to a minimum in order not to deter the more casual The majority of species, except for mycophagous Rhy- user. Fuller bibliographies can be found elsewhere, as sodidae, are predacious. Rhysodidae and Carabidae are indicated above, although for some families very little has terrestrial, Dytiscidae and Gyrinidae are aquatic. been published that might be helpful in a New Zealand context. Morphological structures named in the text are intro- [1] Family RHYSODIDAE duced in the section Morphology and Biology (p. 14); Fig. 101 most are also illustrated. A Glossary (p. 91) defines most Length 5-10 mm Tarsal formula 5-5- 5 morphological terms, along with many others used in this publication that may be unfamiliar. DIAGNOSIS. Body narrowly elongate, subcylindrical Two alternative dichotomous keys are provided for to slightly flattened, widest at middle of elytra, glabrous identification of New Zealand Coleoptera. They represent (except for tactile setae), black or exceptionally reddish slightly different concepts in identification, and are meant brown, strongly glossy. Head grooved and strongly to enable users to confirm an identification by using both. constricted posteriorly to form a narrow neck. Ant- The pyramidal Key I (p. 97) is divided into several ennae stout and beadlike (Fig. 19). Pronotum with 1-3 components, and groups families sharing distinct features deep, longitudinal grooves. Legs short, robust; procoxae regardless of their systematic relationships; it represents globose, their cavities closed. the shorter identification route, and is intended for rapid or preliminary identification. The linear Key II (p. 106) is not SYNOPSIS. There are 6 species of Rhysodidae known subdivided, and represents slightly the longer identification from New Zealand, in 4 genera, of which 2 are endemic.. route. The two keys often employ different character sets. Adults and larvae are thought to feed on plasmodia of They are designed to identify the families of the New Myxomycetes (J.F. Lawrence, pers. comm.), and live in- Zealand coleopteran fauna, and may not be satisfactory for side dead or rotten wood. They may be found under bark or other faunas. Characters which require dissection or dis- in wood of logs, standing dead trunks, or stumps in a partly memberment of specimens have been avoided. Both keys decayed but not excessively wet or dry condition. have been thoroughly tested, but may still contain errors or Collecting: sifting leaf litter, hand picking from under inconsistencies. In the pyramidal key some families with bark, or crumbling and sifting pieces of moderately rotten characters which might be overlooked, or are dubious, or old logs. which differ between the sexes reappear under different Groups. Identifications derived from the keys may be RANGE. North ®, South ®?, Stewart O checked against the appropriate habitus illustration(s) and, Kermadecs O, Three Kings O, Chathams O, Snares O if des-ired, the family diagnosis. Cross-reference to figure Aucklands O, Campbell O, Antipodes O, Bounties O num-bers and family numbers is provided for convenience. For the benefit of anyone using this publication exten- EXAMPLE. Rhyzodiastes proprius (Broun, 1880) (Fig. sively for identifications at the microscope, we strongly 101). recommend that you photocopy either the keys or the illustrations (or both), and use them conjointly flat on the REMARKS. The Rhysodidae are sometimes treated as bench-top. Single copies for this purpose are not in breach tribe Rhysodini of Carabidae. of the publisher's copyright. Suggested colloquial name: heraldic beetles.
-20- SELECTED®, REFERENCES. BellAntipodes & BountiesBell (1978, 1979, Aucklands ®, Campbell 1982, 1985), Bell (1991), Beutel (1990), Emberson (1995), Watt (1980a). EXAMPLES. CICI E I AE - Neocicindela tuberculata (Fabricius, 1775) (Fig. 102); CA A I AE - M r p b r f rb rn r 19 3 (Fig. 1 3 ; MIGA O I AE - x- [ ] l CA A I AE r n br d Guérin-Mén v ll 1 1 (Fig. 104); (incl. CICI E I AE SCA I I AE - Clivina basalis Chaudoir, 1 3 (Fig. 1 5 ; [assisted by R.M. Emberson and J.F. Lawrence] Τ ΕCΗΙΝΑΕ - ll n alacris (Broun, 1 1 (Fig. 106); A A I AE - Ctenognathus n v z l nd Fairmaire, Fig. 102-107 (Fig. 1 3 1 7 Length 1.4-39 mm Tarsal formula 5-5-5 REMARKS. The most recent subfamily classifications of DIAGNOSIS (see Fig. 1, 2). d n r ll r r Carabidae have been proposed by Erwin (1991), Beutel l tr l n d th r f p n t r f f x d (1992, 1993), Arndt (1993), and Lawrence & Newton p t n b r n t t l t usually glossy and black (1995). In older classifications the subfamily Cicindelinae but in some species colourful and spotted. Head usually was regarded as a distinct family. flattened, prognathous, with h rp tr n nd bl Bousquet & Larochelle (1993) proposed a classification f n f r rd Ant nn 11- nt d ll f l - of Carabidae with 17 tribal divisions only, because of the f r n rt d b t n nd b f nd bl lack of consensus among authors at the subfamily and (except Cicindelinae). Eyes prominent. h r x th supertribe level. Dr J.F. Lawrence has provided a key to th l t r l r n ft n ll d f n d El tr pl tribes of Carabidae in New Zealand (Appendix 3); This ll lpt r d th l n t d n l r v (striae). may be preferred by some users over the subfamily key ll tr n t d f r r nn n ; pr t b th below. p l nt nn l l n n r n consisting of an exca- No comprehensive treatment of New Zealand Cara- vation lined with a comb-like setal fringe; pr x ft n bidae is available. However, the group is under review by l b th r v t p n r l d Dr Rowan Emberson (Lincoln University) and by Mr André Larochelle (associate, Landcare Research, Auck- SYNOPSIS. The carabids, one of the largest beetle fam- land), among others. ilies in New Zealand, consist of 6 subfamilies (Lawrence Suggested colloquial name: ground beetles (Cicindelinae & Newton 1995) – CA A I AE CICI E I AE MIGA O I AE – tiger beetles). SCA I I AE EC I AE and A A I AE - with some 445 species in approximately 75 genera and 16 tribes. KEY TO SUBFAMILIES OF CARABIDAE Many carabids are unique to New Zealand, e.g., the Modified from Lawrence & Britton (1994), following the endemic genus Mecodema with some 60 species, mostly arrangement of Lawrence & Newton (1995). For identi- large and robust in form, which apparently occupy similar fication of tribes, consult the key provided as Appendix 3. ecological niches to the northern temperate genus Cara- bus. (Several specimens of Carabus were found in Auck- 1 Antennae inserted dorsally on frons; clypeus broader land in the 1940s, but none is established.) Many carabids than distance between antennal sockets; eyes ex- are apterous, and often occupy restricted habitats or tremely large and protruding; legs very long and geographical areas. There are interesting large species of slender; protibia with 2 terminal spurs; lateral pronotal Megadromus, e.g., M. antarcticus, which is common on carina absent or incomplete posteriorly the Canterbury Plains in the South Island. Small species of ... (Fig. 102) .. CICI E I AE the large genus Bembidion may commonly be found on —Antennae inserted on side of head between eye and banks of streams, rivers, and lakes. mandibular scrobe; clypeus narrower than distance A dominant group of terrestrial predators, with some between antennal sockets; eyes moderately large, not known to be arboreal, mainly nocturnal but some diurnal. or slightly protruding (exception: Scopodes species, Encountered in a variety of habitats ranging from forest which have large protruding eyes); without other com- litter, old tree trunks, under stones, and tussock grass to bination of characters ... 2 some wet habitats, including riparian and coastal areas. Collecting: baited or unbaited pitfall traps, on the 2(1) Metepimeron not visible between posterior edge of ground and around the base of tree trunks, sifting organic metepisternum and anterior edge of 1st ventrite; debris, and hand-picking from logs or under stones or mesocoxal cavities open laterally, partly closed by bark, at night with a head-light. mesepimeron ... 3 —Metepimeron visible as a lobe between metepisternum RANGE. North®, South Stewart ® and 1st ventrite; mesocoxal cavities usually closed Kermadecs ®, Three Kings Snares ®, Chathams ® laterally by meeting of sterna ... 4
- 1- 3(2) Procoxal cavities open behind; apical segment of All our dytiscids are aquatic and very good swimmers. maxillary palp more or less expanded and truncate They often occur in small bodies of water such as ponds, apically; body length more than 17 mm but may be found in streams and rivers, and even under- ... (Fig. 103) .. CARABINAE ground in waterlogged coarse alluvial gravels. They norm- -Procoxal cavities closed behind; apical segment of ally occur in well vegetated waters, where adults and maxillary palp slender and fusiform; body length larvae prey on small arthropods. Dytiscids breathe air usually 17 mm or less ... (Fig. 104) .. MIGADOPINAE trapped under the elytra, and periodically come to the surface to renew their air supply. Most species fly, and may 4(2) Body pedunculate, with a distinct, narrow waist or migrate to different sites. Of special interest are phreatic peduncle between prothorax and elytra, the scutellum water beetles of the genera K h l dr and hr t - entirely contained within peduncle ... 5 d with species living in underground waters (Ordish - Body not pedunculate, or if slightly so then scutellum 1976a, 1991). extending behind peduncle and forming a wedge Collecting: using an aquatic net in small ponds and between elytral bases ... 6 other open waters, or by light trapping near water. Samp- ling the phreatic (underground) fauna requires special 5(4) Mesocoxal cavities open laterally, partly closed by techniques (see, e.g., Ordish 1976a, 1991). mesepimeron ... (Fig. 105) .. SCARITINAE - Mesocoxal cavities closed laterally by meeting of RANGE. North ®, South Stewart sterna ... (Fig. 106) .. TRECHINAE (in part) Kermadecs ®, Three Kings O, Chathams ®, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O 6(4) Mandible with 1 setiferous puncture in a scrobe along outer edge ... (Fig. 106) .. TRECHINAE (in part) EXAMPLES. HYDROPORINAE - K h l dr phr t - Mandible without a setiferous puncture in a scrobe Ordish, 1976 (Fig. 108), d pl t Sharp, 1882 ( ί 1 7 A A I AE (Fig. 109).
SELECTED REFERENCES. Bousquet (1991b), Britton REMARKS. Suggested colloquial name: diving beetles. (1940, 1941, 1949, 1958, 1959, 1960a, b, 1962, 1963, 1964), Brouerius van Nidek (1965), Butcher (1984), SELECTED REFERENCES. Ordish (1966, 1974, 1976a, Gourlay (1950), Johns (1974), Lindroth (1976, 1980), b, 1989, 1991), Spangler (1991a). Moore (1980), Pilgrim (1963), Rivalier (1964), Townsend (1965), Watt (1961), Wise (1988, 1990). [4] Family GYRINIDAE Fig. 110
[3] Family DYTISCIDAE Length 4.3-4.5 mm Tarsal formula 5-5-5 Fig. 108, 109 DIAGNOSIS. Body glossy, boat-shaped, moderately convex to slightly flattened. Eyes completely divided into Length 2.5-27 mm Tarsal formula 5-5-5 upper and lower portions by a strip of cuticle. Antennae DIAGNOSIS. Body elongate-oval, smooth, convex, short, compact, usually clubbed. Middle and hind legs with dorsal and ventral surfaces of similar convexity, short, with fringes of swimming hairs; forelegs pro- dark brown, dark olive, or nearly black, often with nounced and raptorial, with procoxae transverse, and indistinct yellowish spots or margins. Antennae slender their cavities open. and threadlike. Mandibles strong. Scutellum visible or not. Procoxae globose, their cavities open or closed; SYNOPSIS. Represented by one species of GYRININAE, protibiae lacking spines. Hind legs enlarged, modified G r n nv x l self-introduced to New Zealand for swimming (moving synchronously), with paddle- from Australia. First recorded from New Zealand as like tarsi bearing a dense fringe of hairs; metacoxae G r n h tt n (1878), the original specimen was col- large, lacking plates; metatarsi with either 1 or 2 claws. lected by Capt. F.W. Hutton in the Waikato basin in 1873. Sometimes males with first 3 basal segments of protarsi No other specimens were seen until the late 1970s, when dilated to form adhesive pads consisting of suction discs, the species was rediscovered in the Waikato basin. Mr enabling male to hold female during copulation. Ventral Keith Wise (formerly of Auckland Museum) recently body without a flattened keel. found it in the Ahipara area of Northland. Immigrant populations appear to establish in New Zealand SYNOPSIS. Dytiscids are represented in New Zealand sporadically but not widely. by some 16 species in 11 genera of subfamilies COPE- The species occurs in small peaty lakes and man-made LATINAE, HYDROPORINAE, COLYMBETINAE, and DYTISCINAE. dams. Collecting: using aquatic nets.
-22- RANGE. North ®, South ®?, Stewart O cent club; segment preceding club (`cupule') glabrous Kermadecs O, Three Kings O, Chathams O, Snares O and often embracing the following segment. Maxillary Aucklands O, CampbelI 0, Antipodes O, Bounties O palps elongate, often longer than antennae. Legs often dentate or spinose; procoxae large, transverse and pro- EXAMPLE. GYRININAE - Gyrinus convexiusculus Mac- jecting, their cavities open; metatarsi flattened and leay, 1871 (Fig. 110). fringed with hairs. Metasternum often extended at rear into 4 short spines. REMARKS. Suggested colloquial name: whirligig beetles. SYNOPSIS. Approximately 75 species of hydrophilid SELECTED REFERENCES. Ochs (1949), Spangler are recorded in New Zealand, in some 25 genera and 3 (1991b), Wise (1989). subfamilies - HORELOPHINAE, HYDROPHILINAE, and SPHAER IDIINAE. Horelophinae are endemic to New Zealand; the type genus Horelophus is a very atypical hydrophilid. Suborder POLYPHAGA There are two main groups of species, aquatic and Comprises the vast majority of Coleoptera-95% ac- terrestrial; many New Zealand hydrophilids are terrestrial, cording to Lawrence & Britton (1991, 1994), whose and are found in leaf litter or carrion. Larvae tend to be definition we follow here: prothoracic notopleural suture aquatic or semiaquatic, but many species are known from absent (Fig. 3); prothoracic pleuron fused with trochantin moist, decaying organic matter. A few species are reported and entirely concealed, forming a cryptopleuron; hind from thermal waters. Adults are usually herbivorous or wing without an oblongum cell, and with transverse fold scavengers, whereas all known larvae are carnivorous and never crossing MP (Fig. 51); 1st abdominal sternite not cannibalistic. divided by metacoxae (Fig. 3), which are usually movable Collecting: netting in aquatic habitats, sifting decaying and not fused to metasternum; aedeagus usually with a vegetable matter, or light-trapping. basal piece. RANGE. North ®,® South . Stewart ® Kermadecs. Three ® Kings ®, Chathams ®, Snares ® Series STAPHYLINIFORMIA Aucklands ®, Campbell O, Antipodes ®, Bounties ® Adults distinguished by reduced wing venation (except in Hydrophilidae) and a sophisticated type of wing-folding EXAMPLES. O E O I AE - Horelophus walkeri Orch- mechanism; larvae usually with articulated urogomphi ymont, 1913 (Fig. 111); ΗY OΡΗI ΙΝΑΕ- Enochrus tritus (Lawrence & Britton 1991, 1994). (Broun, 1880) (Fig. 112); SΡΗΑΕ I ΙΙΝΑE - Rygmodus tibialis Broun, 1886 (Fig. 113).
Superfamily HYDROPHILOIDEA REMARKS, New Zealand's diverse fauna of Hydro- Antennae short, with a long scape and a densely pubescent philidae is much in need of revision at all levels. The 3-segmented club (Fig. 26, 27); segment preceding club Sphaeridiinae are under review by Dr M. Hansen (Zoo- (`cupule') transverse, concave, and in aquatic species used logical Museum, Copenhagen). The terrestrial forms are for respiration. Hind wings with a radio-median (R-M) probably important indicators of environmental quality. loop, without a spring mechanism. Procoxae large; tibiae Suggested colloquial name: water scavenger beetles. dentate or spinose; tarsi 5-5-5, 5-5-4, or 5-4-4, sometimes appearing 4-4-4 because of vestigial basal segment. Aede- SELECTED REFERENCES. Hansen (1991b, 1995), agus trilobed (with fused parameres in most Histeridae). Newton (1989), Orchymont (1913), Ordish (1974, Larva distinct (for details see Lawrence & Britton 1991, 1976b), Spangler (1991d), Todd (1961), Winterbourn 1994). A phylogenetic analysis of Hydrophiloidea based (1968, 1970), Wise (1965, 1973). on characters of the adult head and the larvae is provided by Beutel (1994) and Hansen (1991a, b, 1995). [6] Family HISTERIDAE Fig. 114, 115 [5] Family HYDROPHILIDAE Length 1.5-10 mm Tarsal formula 5-5-5 or 5-5-4 Fig. 111-113 Length 2-9 mm Tarsal formula 4-4-4 or 5-5-5 DIAGNOSIS. Body robust, disc-shaped, oval or nar- rowly oval to nearly rectangular with rounded angles, DIAGNOSIS. Body usually oval, moderately to glabrous, moderately convex to flattened, black and strongly convex, smooth and usually glossy. Antennae strongly glossy. Head deeply inserted. Antennae unique short, 7-9-segmented, often concealed from above, in form: short and elbowed, with an abrupt, 3- with a long scape and a 3-segmented, densely pubes- segmented club, often inserted in a cavity or hidden on
-23- underside of prothorax. El tr h rt n d nd tr n t [7] l Y AE I AE exposing tip of abdomen (usually 2 or 3 apical segments), Fig. 116, 117 with 6 or fewer striae. r x l r nd tr n v r th r v t p n; pr t b xp nd d ft n p n Length 1.3-2.6 mm Tarsal formula 5-5-5 DIAGNOSIS. d n rr l v l r ll pt l (broad SYNOPSIS. There are 29 nominal New Zealand species in species of Meropathus), flattened to slightly convex, of histerid, of which 6 are adventive. They occupy some 13 small, with scutellum small; generally similar to Hydro- genera, including one undescribed, in 5 subfamilies - philidae but th r 7 bd n l nt pp d A AEI AE SA I I AÉ E O I I AE I A I AE and t 5 M x ll r p lp l n t ll l n r th n IS E I AE Parepierus is the largest genus, containing nt nn Ant nn 9-11- nt d (appearing 8- about one-third of the species, and Reichardtia is an segmented in Meropathus) and th - 3- r ll 5- endemic New Zealand genus. Most species are widely nt d p b nt l b th l t t r r distributed, although histerids are infrequently encountered l n t often longer than all preceding segments com- in New Zealand. bined, with no distinct bisetose empodium between claws; Adults and larvae are almost exclusively carnivorous, pr x tr n v r th r v t p n r (Hydraena) and prey on other insects, usually larvae. They may be l d commonly found on carrion, decomposing plant materials, tree wounds, seashore debris, rotting mushrooms, and SYNOPSIS. Thirty-two species of hydraenid are recog- under bark in galleries of other insects. Kuschel (1990) nised in New Zealand, in 5 genera and 2 subfamilies - recorded some species in a suburban environment from Y AE I AE and OC E II AE Twenty-two species are compost, lawn clippings, poultry straw, carcasses, fowl recorded from the South Island and subantarctic islands, 4 manure, bush floor litter, and decayed wood. The smaller from the North Island, and 6 occur on both main islands species are sometimes common around seabird nests. (Ordish 1984). This relatively rich hydraenid fauna offers Collecting: hand-picking, or sifting organic material an interesting field for ecological studies. and processing it in Berlese funnels. All the genera except Meropathus live as adults and larvae in fast-flowing streams. Meropathus species are RANGE. North ®, South ®, Stewart terrestrial in the coastal or supralittoral zone, occurring in Kermadecs ®, Three Kings ®, Chathams ®, Snares O porous rocks, littoral moss and grasses, and seabird nests. Aucklands O Campbell O Antipodes O Bounties O Adults of the aquatic genera all have one or more pairs of legs modified (Ordish 1984, fig. 12-16) and feed on dead EXAMPLES. SAPRININAE - Reichardtia pedatrix (Sharp, leaves. 1876) (Fig. 114), Tomogenius latipes (Broun, 1881) (Fig. 115). RANGE. North ®, South ®, Stewart Kermadecs O Three Kings O Chathams . Snares ® REMARKS. This family is under revision by Dr Masa- Aucklands ®, Campbell ® Antipodes® ®, Bounties O hiro Ohara of Hokkaido University, Japan. Suggested colloquial name: hister beetles, pill beetles. EXAMPLES. HYDRAENINAE - Podaena latipalpis Ordish, 1984 (Fig. 116); OCHTHEBIINAE - Meropathus zelandicus SELECTED REFERENCES. Dahlgren (1968, 1971, Ordish, 1984 (Fig. 117). 1976), Mazur (1984), Newton (1991 c), Wenzel (1944). REMARKS. Suggested colloquial names: minute moss beetles, cascade beetles. S p rf l S A Y I OI EA Body shape variable, antlike in Scydmaenidae (Fig. 122), SELECTED REFERENCES. Gressitt & Samuelson short, stout, and wedge-shaped in Pselaphinae (Fig. 126), (1964a), Hansen (1991a), Ordish (1984), Spangler ovoid in Hydraenidae (Fig. 116, 117) and Agyrtidae (Fig. (1991c).
119 ; ovoid to nearly spherical in Ptiliidae and Leiodidae (Fig. 118, 120, 1 1 and elongate, slender, with exposed flexible abdomen in most Staphylinidae (e.g., Fig. 132- [ ] l I II AE 138). Adults distinctive in having usually strongly Fig. 118 projecting coxae, metasternum almost always lacking a Body length 0.6-1 mm Tarsal formula 2-2-2 or 3-3-3 median suture, legs often spinose, elytra usually truncate, exposing at least one tergite, hind wing with reduced DIAGNOSIS. d n t brown to black, broadly venation and without a radio-medial (R—M) loop, and elongate-oval. Ant nn f l nt b r n l n aedeagus with a reduced phallobase. Larvae characteristic h r nd th - r 3- nt d l b El tr ft n — see Lawrence & Britton (1991, 1994). h rt xp n 1 r bd n l nt nd n
— — f th r-l fr n d th l n h r Scutellum large. (1982), and are believed to be closely related to Leiodidae r x tr n v r r l b th r v t p n; (Newton 1985). metacoxae variable in shape, often with large coxal plates. Species of Agyrtidae are known from forested areas, Tarsi usually 2- or 3-segmented but appearing 1-segmen- and are common on carrion and faeces. Adults of `N'. ted in some species. prolongatus occur in forested areas on carrion, e.g., dead birds especially at a slightly mummified stage, or may be SYNOPSIS. In New Zealand there are some 56 species of found in soil crevices on stream banks. Some specimens ptiliid, including 8 or 9 adventive, in perhaps 13 genera were collected from an abandoned kiwi egg in a burrow. and 3 subfamilies — PTILIINAE, NANOSELLINAE, and ACRO- An effective method of collecting adults is carrion-baited TRICHINAE. pitfall traps. Larvae may be seen in association with adults Common in decaying organic matter, where apparently on carrion. they feed on fungal spores and hyphae (Lawrence & Britton 1991, 1994). Collected from garden compost, leaf RANGE. North ®, South ®, Stewart O litter, moss mat, forest litter, decayed wood, mouldy dung, Kermadecs O Three Kings O Chathams O Snares O under bark in native forests, amongst litter and tussocks, Aucklands O Campbell O Antipodes O Bounties O seashore debris including decaying seaweed, and humi- fied soil (Johnson 1982, Kuschel 1990). Adults are often EXAMPLE. `Necrophilus' prolongatus (Fig. 119). caught in flight interception traps. REMARKS. Suggested colloquial name: carrion beetles. RANGE. North . South ®, Stewart ® Kermadecs . Three Kings ®,® Chathams ® Snares ® SELECTED REFERENCES. Newton (1991a). Aucklands ®, Campbell ®, Antipodes ® Bounties O
EXAMPLE. PTILIINAE - Notoptenidium lawsoni (Mat- [1 ] l EIO I AE thews, 1873) (Fig. 118). [assisted by S.B. Peck and A.F. Newton, Jr] Fig. 120, 121 REMARKS. Suggested colloquial name: feather-winged beetles. Body length 0.9-4.3 mm Tarsal formula 5-5-5, 5-5-4, or 5-4-4 SELECTED REFERENCES. Dybas (1991), Gressitt & DIAGNOSIS. d br dl v l t n rl ph r l Samuelson (1964a), Johnson (l975a—c, 1982). l htl t tr n l nv x r d n d-l ; rf l br nd tr n l l r p b nt brown to black. Ant nn th 3-5- nt d l b; h n l b [9] l AGY I AE 5- nt d th n nt ( nd nt f l b Fig. 119 ll ll r th n dj n n nt (Fig. 24);
Length 9-10 mm Tarsal formula 5-5-5 d t l rf f nt nn l nt 7 9 nd 1 th p n n f n t p f n r v l Elytra often DIAGNOSIS. d br dl v l l htl d pr d striate, and often with a longitudinally impressed line th d f pr n t nd l tr xpl n t brown, along suture. r x l b nd pr j t n ; t - glabrous and glossy. Ant nn 11- nt d th 5- x l t th r b ft n xp nd d nd p n nt d p b nt l b El tr v r n nt r bd n th 9 r 1 tr Hind wings well dev- SYNOPSIS. According to unpublished data of G. eloped. r x n l l htl tr n v r b ll Kuschel, A. Newton, and S. Peck (pers. comm.) there are nd pr j t n th r v t p n Mesotibiae and some 112 species in approximately 25 currently recog- metatibiae each with a large spur. r r nd l nised genera, including 6 or 7 new genera and nearly 50 l r new species. Four subfamilies are represented — CAM'- ARINAE, with some 69 species, is the largest, followed by SYNOPSIS. A small, archaic family with fossil records CHOLEVINAE (25), LEIODINAE (17), and COLONINAE (1). from the mid Jurassic (Newton 1991a), comprising Kuschel (1990) reported collecting species of Leio- approximately 8 genera in the world fauna. Most agyrtids dinae in the Auckland area from bush-floor litter, amongst are Holarctic in distribution, with the exception of two ground plants, from grass mats of Oplismenus and New Zealand species: ` Necrophilus' prolongatus Sharp, Microlaena, and from an agaric in the bush. He recorded and an undescribed species (Newton 1985). According to Camiarinae collected in the same area from fungi (Auri- Newton these belong in an undescribed genus, and are cularia polytricha, Ganoderma applanatum), coarse floor erroneously placed in Necrophilus. Agyrtidae were re- litter, amongst Gahnia plants, under logs and in cracks of moved from family Silphidae by Lawrence & Newton stream banks, decayed wood, in Ptychomnion moss mats,
— 5 — and rotten n r d t and Kn ht logs. He collected SELECTED REFERENCES. Daffner (1985), Newton Coloninae in bush litter, cracked stream banks, and (1991b), Szymczakowski (1964, 1966, 1973). Malaise traps, and collected Cholevinae from carrion. Newton (1984) reported adults and larvae f r t d (Camiarinae) collected in numbers on bracket fungi, with [11] Family SCYDMAENIDAE the following new host records: l f r and several [assisted by G. Kuschel] allied undescribed species on G n d r t p r Fig. 122 G ppl n t and unspecified bracket fungi, all from the North Island. Adults and larvae were browsing on the Length 0.5-3.5 mm Tarsal formula 5-5-5 hymenial surface of G n d r Gut contents of both DIAGNOSIS. Body reddish brown to dark brown, usu- stages included spores and hyphae. Newton (1984) ally ant-like, with a distinct `waist' constriction be- provided some data for the worldwide genus C l n as tween prothorax and elytra. Head with a distinct neck. collected in flight traps and by sweeping vegetation, Antennae long, pubescent, with a more or less distinct especially grasses in wooded areas, and suspects that the 3- or 4-segmented club. Pronotum ovoid or spherical, genus is associated with hypogeal fungi or moulds. often with basal foveae, and without carinae. Elytra Species of Cholevinae, both adults and larvae, may be oval, covering entire abdomen, with rounded humeri; found on decaying Organic matter, on which they feed: entire dorsal surface usually glossy and with long hairs. dung, carrion, well decayed soft fungi, forest litter, nests of Maxillary palps usually enlarged, with apical segment vertebrates, etc. (Newton 1984). reduced. Legs with enlarged femora. Procoxae trans- verse and projecting, their cavities open. RANGE. North ®, South ®, Stewart O, Three Kings ®, Chathams ®, Snares Kermadecs SYNOPSIS. Watt (1982a) recorded 165 nominal native Aucklands ®, Campbell ®, Antipodes O, Bounties O species of scydmaenid; 201 native and 1 adventive species are now represented in NZAC, many of them undescribed. EXAMPLES. CAMIARINAE - In t p l n ll Broun, They belong to some 11 currently recognised genera of 1917 (Fig. 120); COLONINAE- C l n h rt l (Broun, 1880) SCYDMAENINAE, of which 6 are endemic to New Zealand. (Fig. 121). For classification see Franz (1975, 1977, 1980, 1985). Kuschel (1990) recorded several species from the REMARKS. Suggested colloquial name: small carrion Auckland area collected in leaf litter, from decayed wood beetles. of various trees and hollow logs, moss mat, leaf mould in tree hollows, and Malaise traps. Scydmaenids are noc- KEY TO SUBFAMILIES OF LEIODIDAE turnal, living in concealed habitats, with some species IN NEW ZEALAND [by A.F. Newton, Jr] flying at dusk. Good collections might be obtained by 1 Head with an elevated crest (occipital carina) along processing sifting organic litter through Berlese funnels. hind margin, resting against front of pronotum when head in repose ... CHOLEVINAE RANGE. North ®, South ®, Stewart ® —Head without an elevated crest along hind margin .. 2 Kermadecs O, Three Kings ®, Chathams ®, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O 2(1) Antennae 11-segmented, with segment 8 as large as segments 9 and 10, and club of 4 segments (Fig. 25); EXAMPLE. SCYDMAENINAE - Adr t l r (Franz, dorsum setose ... (Fig. 121) .. CO O I AE 1985) (Fig. 122). —Antennae usually 11-segmented, with segment 8 smaller than segments 7 and 9, and club usually of 5 REMARKS. Suggested colloquial name: stone beetles. segments (Fig. 24); if club 4-segmented then antennae 10-segmented and dorsum glabrous ... 3 SELECTED REFERENCES. Franz (1975, 1977, 1980, 1985). 3(2) Antennal insertions concealed in dorsal view; head relatively flattened and broad, usually half or more as wide as pronotum; dorsum apparently glabrous (sometimes with many short hairs visible only at [12] Family STAPHYLINIDAE (incl. PSELAPHINAE, SCAPHIDIINAE) higher magnification) ... EIO I AE —Antennal insertions exposed in dorsal view; head [assisted by A.F. Newton and M.K. Thayer] relatively convex and narrow, usually less than half as Fig. 123-138 wide as pronotum; dorsum usually with evident long
setae (exceptions: A ph r t C t p l Ch l - Length 0.6-25 mm Tarsal formula 5-5-5 rt d (Fig. 120) .. CAMIA I AE or heteromerous
- - DIAGNOSIS. Body brown to black, less frequently vegetation. The majority of known species are predators, brightly coloured or metallic. Form very diverse, usually feeding on larvae and adults of other insects including narrowly elongate, approximately subparallel, rarely Collembola, and on mites and nematodes. short, oblong, or wedge-shaped, or short, stout, and oval Species of Osoriinae and Oxytelinae feed on decom- with abdomen non-flexible (Pselaphinae, Fig. 126). Head posing organic matter (McColl 1982, Newton 1990). often with a posterior constriction, sometimes with ocelli Some Tachyporinae and Aleocharinae (e.g., G r ph n (Omaliinae, Microsilphinae). Antennae with 10 or 11 feed on moulds or other fungi. Species of the aleocharine segments, filiform, beadlike; last 1-4 segments enlarged genus M ll n were recorded feeding on algae (Klima- (Pselaphinae), incrassate, or clubbed. Elytra nearly szewski 1992). Larvae of the ectoparasitic genus Al - always short and truncate, exposing a variable number h r (Aleocharinae) are exclusive parasitoids of puparia of abdominal segments, usually 6 or 7, exceptionally of cyclorrhaphous Diptera. The first-instar larva searches concealing alI or most of abdomen (Microsilphinae and for a host pupa, gnaws a hole in the puparium, crawls some Omaliinae, Proteininae, Scaphiidinae) or leaving at inside, seals the entrance behind, and feeds on the pupa least 2 (usually 3 or 4) tergites exposed (Pselaphinae). (Klimaszewski & Crosby 1997; Klimaszewski & Jansen Procoxae usually conical and strongly projecting, their 1993). cavities usually closed. Abdomen flexible dorsoventrally The majority of adult pselaphines occur in leaf and (exception: Pselaphinae), well sclerotised except basal wood litter on the forest floor. Newton & Chandler (1989) tergite or two. reported that in general pselaphine species can also be discovered in wetland, grassland, desert, beach, cave, and SYNOPSIS. Staphylinids are one of the largest and most even arboreal habitats, wherever debris, moss, or root mats diverse families of beetles, as of 1987 comprising world- maintain microhabitats of high humidity. Kuschel (1990) wide about 42 000 nominal species (8400 species of collected pselaphines in Auckland from decayed wood on Pselaphinae) in several thousand genera (1091 genera in the beach and amongst low coastal vegetation, coastal Pselaphinae) and 100 tribes (Newton 1985, 1990, Newton litter, hollow trees, wood mould, stream litter and stream & Chandler 1989, Newton & Thayer 1992). Hundreds of banks, moss, sedges, grasses, soil around tree stumps, etc. new species are described every year. We follow Newton Adults and larvae of pselaphines are predators, feeding on & Thayer (1995) and Lawrence & Newton (1995) in mites, springtails, and other small invertebrates. treating the former family Pselaphidae as a subfamily of More details on the natural history of New Zealand Staphylinidae. A revised list of the subfamilies, tribes, and staphylinids are provided by Klimaszewski t l (1996). genera with synonyms is provided by Klimaszewski t l Collecting: hand-collecting with the aid of an aspirator; (1996). pitfall traps, unbaited or baited with carrion, faeces, or In New Zealand there are approximately 766 nominal fermented fruits; processing sifted organic litter through native species and 85 adventive, plus 170 or more recog- Berlese funnels or Winkler/Moczarski extractors; Malaise nised but not yet named. The number of New Zealand traps, window traps, and other flight interception traps. native staphylinids may exceed 1000 (G. Kuschel, pers. comm.), making it after Curculionidae our second largest RANGE. North ®, South ®, Stewart group of beetles. Our species are grouped in some 190 Kermadecs ®, Three Kings ®, Chathams ®, Snares ® genera and 16 subfamilies, as follows: A EOC A I AE 61 Aucklands ® C pb ll ® Ant p d ® nt Ο genera (probably an underestimate); EUAES E I AE 4; A OCE I AE 1; MIC OSI I AE 1; OMA II AE 16, and 4 EXAMPLES. ΜIC OSI I AE — Microsilpha l t r undescribed; OSO II AE 3 (including one of former Broun, 1 (Fig. 123); OMALIINAE — Omaliomimus Eleusiinae); O Y E I AE 7; AE E I AE 1 ; OEOC A I AE albipennis (Kiesenwetter, 1 77 (Fig. 1 ; Ρ ΟΤΕΙΝΙΝΑE 2; IES I AE 1; O EI I AE 4; SE A I AE 41, and 20 nitidus Broun, 1 95 (Fig. 1 5 ; PSELAPHINAE —-S lph t l undescribed; SEU O SI AE 1; SCAPHIDIINAE 5; S A Y I I AE S l laminata Broun, 1 93 (Fig. 1 ; Ρ OEOC A I AE 26; and AC Y O I AE 3 (Klimaszewski t l 1996; Watt — Pseudophloeocharis australis (Fauvel, 19 (Fig. & McColl n McColl 1982, updated). 1 7 ; ΤAC Y O I AE — Sepedophilus sp. (Fig. 1 ;
Staphylinid beetles are very successful and broadly HABROCERINAE — Habrocerus capillaricornis (Gravenhorst, distributed, occurring in almost all terrestrial habitats. 1 (Fig. 1 9 ; Α EOC A I ΑE —Aleocharah nd Staphylinids may be found in decomposing organic matter (Klimaszewski Cr b 1997 (Fig. 130); SCAPHIDIINAE such as garden and forest litter, mushrooms, bird and — ` r t ll r (Redtenbacher, 1 7 (Fig. 131 ; mammal nests, decaying seaweeds and other algae of tidal ΡIΕS IΝΑΕ—19 (Cameron,(Fig. r n h d n zones, uncompacted soil, organic matter in tree hollows 13 ; OSORIINAE — t t r h ferrugineus (Broun, 1893) and burrows, under bark, in galleries of other insects, wet (Fig. 133 ; OXYTELINAE — Carpelimus sp. (Fig. 134); moss, cracks on edges of streams, shores of lakes, etc. ΕUΑΕSΤΗΕΤΙΝΑΕ — Agnosthaetus v n (Broun, 19 1 Some species may be found on carrion and in animal (Fig. 135 ; ΡSEU O SI AE — Pseudopsis arrowi Bernhauer, droppings, and some Omaliinae have been beaten from 1939 (Fig. 13 ; PAEDERINAE — Medon zeelandicus
- 7- ( dt nb h r 1867) ( 137); S A Y I I AE — Cafius elytra, with head and pronotum narrower; setose litoreus r n 1880 ( 138). foveae present on head, prothorax, and other parts of body; elytra short, closely applied to body, leaving REMARKS. Staphylinids are an ancient group with fossil much of abdomen exposed; abdomen broad, not records from the early Jurassic period (Newton 1990). The flexible; maxillary palps often modified; antennae New Zealand fauna of staphylinids may be characterised often beadlike; tarsi 3-segmented by (1) the absence of several subfamilies from the native ... (Fig. 126) .. SE A I AE fauna, a few of which are, however, represented by ad- —Body not so; setose foveae absent; elytra usually ventive species (e.g., Habrocerinae); (2) large species loosely applied to body; abdomen flexible; maxillary radiations in Eupines, Euplectopsis, Hyperomma, Micro- palps usually simple; antennae not beadlike; tarsi silpha, Ocalea complex, Oligota, Othius, Paratorchus, usually of more than 3 segments ... 5 `Quedius', Sagola, and Sepedophilus; and (3) high endemism — approx. 43% at the generic level, even higher 5(4) Abdomen without paratergites; tergites and sternites at the species level (Klimaszewski et al. 1996); (4) strong usually fused together; body either cylindrical or southern connections — over 20 groups or genera are strongly flattened ... (Fig. 133) .. OSO II AE shared only with other southern temperate regions (New- —Abdomen with paratergites; tergites and sternites of ton 1985). In general our staphylinids are poorly known; abdomen not fused together; body not cylindrical .. 6 the Aleocharinae, as everywhere, are the least known amongst staphylinids, and consequently are in urgent need 6(5) Head with paired ocelli on vertex between posterior of revision. The classification of Staphylinidae at all margins of eyes; paratergites single levels, including family limits, is unsatisfactory and un- ... (Fig. 124) .. OMA II AE stable (Newton 1990). —Head without ocelli (sometimes with a pair of de- Suggested colloquial name: rove beetles. pressions on vertex); paratergites usually double ... 7 7(6) Antennal insertions located between eyes and KEY TO SUBFAMILIES OF STAPHYLINIDAE posterior to a line drawn between anterior edges of OCCURRING IN NEW ZEALAND eyes (except in forms with reduced eyes); terminal [Modified from Watt & McColl in McColl (1982) to segment of maxillary palp usually very small and include Microsilphinae, Pselaphinae, and Scaphidiinae.] sharp-pointed ... (Fig. 130) .. Α ΕΟCΗA ΙΝΑE —Antennal insertions located anterior to a line drawn 1 Antennae distinctly clubbed (e.g., Fig. 123, 135); body between anterior edges of eyes, or inserted under without setose foveae ... 2 shelf-like dorsolateral margins of frons; terminal —Antennae usually not distinctly clubbed; if so, body segment of maxillary palp usually not as above ... 8 compact and with setose foveae, and tarsi 3-segmented (e.g., Fig. 124-134) ... 3 8(7) Body strongly depressed; antennae inserted under widely explanate dorsolateral projections of frons, 2(1) Antennal club with more than 2 segments; elytra their sockets facing ventrally; procoxae globular elongate, covering most of abdomen; anterior margin ... (Fig. 132) .. IES I AE of labrum not as below; body short, oval; head with —Body rarely strongly depressed; antennae usually not paired ocelli on vertex... (Fig. 123) .. MIC OSI I AE as above, but if inserted under dorsolateral projections —Antennae with a 2-segmented club; elytra short, ex- of frons then these are less prominent, and their posing entire abdomen; anterior margin of labrum sockets face laterally; procoxae elongate, transverse, minutely crenulate or denticulate; body narrow, elon- or projecting ... 9 gate; head without ocelli.. (Fig. 135) .. EUAES E I AE 9(8) Abdomen usually strongly flattened and broad, with 7 3(1 Body wedge-shaped, short, convex and strongly visible sternites, excluding genital segment (exception: glossy; elytra truncate posteriorly, concealing all but 1 Coprostygnus, with 6 obvious sternites but also with or 2 abdominal tergites, and exposing pointed abdo- elytral striae, which are rare in taxa keying to the men; legs and antennae long and slender alternative, below); gular sutures confluent, or at least ... (Fig. 131) .. SCA I II AE partially so ... (Fig. 134) .. O Y E I AE —Body not so; elytra exposing at least 3 abdominal —Abdomen usually less flattened and moderately broad, tergites (exception: Silphotelus, Proteininae, Fig. with 6 visible sternites (Fig. 124, 127) excluding 125); legs and antennae usually moderately elongate genital segment; gular sutures separate ... 10 and stout ... 4 1 (9 Elytra elongate, leaving no more than 5 abdominal 4(2) Body compact, usually widest at abdomen or tip of tergites exposed (in Silphotelus elytra almost com-
—28— pletely covering abdomen); lateral margin of frons S r SCA A AEI O MIA with a small, semicircular emargination on either side, S p rf l SCA A AEOI EA just in front of eye; body length usually less than 2 mm Characterised by a highly modified prothorax with large (maximum 4 mm) ... (Fig. 125) .. O EI I AE -Elytra shorter, leaving at least 6 abdominal tergites coxae, usually dentate tibiae with a single apical spur, exposed; lateral margin of frons not so ... 11 lamellate antennal club (Fig. 30), no metacoxal plates, 2nd abdominal sternite represented by a lateral portion only, 11(10) Abdominal tergites except the last each with a tergite 8 forming a pygidium, hind wing with reduced diagonal impressed line on either side; pronotum and venation and with a spring mechanism for folding the elytra longitudinally costate; tarsi short, about one- wing, and larvae grub-like and usually C-shaped (Fig. 98) third as long as tibiae; body length 3 mm (Lawrence & Britton 1991). ... (Fig. 136) .. SEU O SI AE -Abdominal tergites without such lines; pronotum and elytra not longitudinally costate; tarsi usually more [13] l UCA I AE than one-third as long as tibiae ... 12 [assisted by B.A. Holloway] Fig. 139-143 12(11) Head with sides converging evenly posteriorly, Length 6-30 mm Tarsal formula 5-5-5 without a neck constriction clearly visible from above (Fig. 127-129) ... 13 DIAGNOSIS. d r b t medium-sized to large, - Head constricted behind eyes to form a distinct neck, brown to black, often with paler spots forming a pattern, clearly visible from above (Fig. 137, 138) ... 15 th r l br nd l r l th d th l d pr n th th ll d v l p d nd bl h h r 13(12) Elytra with epipleura not delimited, their sides ft n x ll d rph - pr n n d n l and completely rounded off; body shape distinctive, as bearing teeth or larger processes. Ant nn d t n t 1 - figured ... (Fig. 127) .. OEOC A I AE nt d ll lb d th b-l l b - Elytra with epipleura clearly delimited by a carina.. 14 n t n f 3 r th l ll t nt th t nn t b h ld t th r (Fig. 30). Pronotum usually 14(13) Antennae stouter, expanded towards apex; head short. Abd n th 5 v ntr t and pronotum usually densely pubescent; body shape distinctive, as figured ... (Fig. 128) .. AC Y O I AE SYNOPSIS. Holloway (1961, 1962, 1963b) recognised -Antennae very slender, filiform (Fig. 18), with long 26 species in 4 genera. There are also 3 adventive species tactile setae; head and pronotum glabrous except for from Australia, each in a separate genus (Holloway 1997). some long tactile setae; body shape distinctive, as Four subfamilies are represented - AESA I AE SY ESI AE figured ... (Fig. 129) .. A OCE I AE AM IMI AE and UCA I AE The species are approximately evenly divided between 15(12) Maxillary palp with apical segment always re- the North and South islands, with two endemic species duced, less than half as long as penultimate segment; restricted to the Chatham Islands and one to the Moko- antennal insertion obscured by a dorsolateral shelf- hinau Islands (Holloway 1961). Approximately half the like expansion of margin of frons species are flightless and half fully winged. The flightless ... (Fig. 137) .. AE E I AE species are often allopatric, with restricted ranges in - Maxillary palp with apical segment variable in length, lowland areas, whereas fully winged species tend to have but at least half as long as penultimate segment; broad and overlapping ranges, though some are restricted antennal insertions exposed to montane areas (Holloway 1963a). ... (Fig. 138) .. S A Y I I AE Hudson (1934) reported lucanids as being mainly noc- turnal, the adults feeding on tree sap, and larvae living on SELECTED REFERENCES. Bernhauer (1939, 1941, old decaying trees or their roots, and with a life cycle of 1943), Cameron (1944, 1945, 1946, 1947, 1950), Fauvel several years. Holloway (1963a) suspects that all adult (1900), Frank (1991), Hammond (1975), Herman (1975), lucanids feed on sap, and are not plant-specific, the larvae Kasule (1966, 1968, 1970), Klimaszewski (1979), Klima- completing their development in rotten wood or soil. It is szewski & Crosby (1997), Klimaszewski t l (1996), widely accepted that the larvae feed on rotten wood or Leschen & Löbl (1995), McColl (1982, 1984a, b), Newton other decomposing organic material. C r t n th p - (1984, 1985, 1989, 1990), Newton & Chandler (1989), l f r is almost certainly myrmecophilous, found in Newton & Thayer (1992, 1995), Park & Pearce (1962), endemic formicine ant nests (Holloway 1962). Hudson Sharp (1876a, b), Steel (1949, 1950a, b, 1953, 1960, 1964, (1934) reported C r t n th bb as being beaten 1966, 1970), Thayer (1985), Williams (1976). from shrubs in hot sunshine, from December until March.
- 9- A GE rth ® S th ® St rt ® [15j[15] l SCA A AEI AE K r d ® hr K n ® Ch th ® ® Sn r O [ t d b M E b r n] A l nd O C pb ll O Ant p d O nt O 1 5-15 n th 3- 5 r l f r l 5-5-5 E AM ES AESA I AE - Ceratognathus parrianus (W t d 1 3 ( 139 ; UCA I AE - Paralissotes reticulatus (W t d 1 7 l ( 1 Geodorcus IAG OSIS St t-b d d b tl ft n l b t Saprosites, auriculatus ( r n 19 3 l nd f l ( 1 1 d ll ( Aph d n l br r d r t l t d n l p b nt ll br n r bl 1 ; AM IMI AE - Dendroblax earlii (Wh t 1 ( 1 3 b t n ll br htl l r d ( M l l nth n E th r l br r nd bl r b th n l d fr b v b l p Ant nn -1 - nt d n t EMA KS h f l nd r r v n b r A lb d; l b v r bl ll 3- - r 5- nt d ll f A l nd b t n t p l f r ; l ll t nt pp r n S t d ll l n t b tl l n t r v l l b th t n b f ld d t th r t htl t f r p t nd tr l l b SE EC E E E E CES C rl n (1991 ll S lpt r f l tr n t r h tr n l d f d f r (19 19 1 19 19 3 b 199 1997 d n th t th p n nd/ r br tl Abd n th v ntr t
[1 ] l OGI AE SY O SIS W tt (19 r rd d ppr x t l 13 1 n t v nd 1 dv nt v p f r b d n l nd h r l t v l l r r p f b tl r pr nt n th - 5 r l f r l 5-5-5 bf l —ME O O I AE SCA A AEI AE ΥΝΑSΤΙΝΑE A O II AE— nd 5 n r 13 f th nd IAG OSIS d r b t v l l htl bp r ll l h bf l M l l nth n b f r th l r t th h v l l r t d th d r l rf r l - 1 3 v l d n t v p n 1 n r (G v n 195 195 tr nd ft n v r d th d br d tr n l 19 ( t n r 19 W tt 19 E r n rr tt d fl x d Ant nn 1 - nt d lb d th 1997 Sp f Prodontria r ll fl htl h v p t l ll t l b ( 31 r x pr j t n ; l t d r n nd r v ln r bl t h b t t d f t n x l v t br dl l d h Cr ll h f r P. lewisi, n d r d t b n nd n r d p nd n t r r rv h b n t- SY O SIS l nd h nl n l dv nt v bl h d n r Cr ll t pr rv t (W tt 19 h E r p n p f tr d h f r t f nd t K nd n b tl Pyronota festiva, n ll n rth- t f A l nd n d p l tt r f p ltr h n v r l nd nd f r t ll t d d r n C pt n t n th f d r t r Oth r r rd C v n 17 9 h 9 r n h- l r d p n l d C pb ll h n r M t n W t rn ll f Stethaspis r n n f r t h b t t h b- n r Wh n r th I l nd nd t n f l nt n n p h h r r n d p t Ad lt h v b n ll t d n r r t r nd r th f r lt r th r r b Costelytra zealandica. h d f l n -d d n dr n l r n n d h S r b n th 1 p n l d n t v ld n nd l nd n b rd n t ( r n 19 7 p n nd n r nd dv nt v p n 3 rv f th r p r n n t r n v rt l x t n r (E b r n M tth 1973 l n 1935 b rr b n th r (C rl n 1991b r n W tt 19 h n t n n l d 5 n t v p r tt n 1991 199 n th nd n Pericoptus (Endröd 197 W tt 19 nd 3 dv nt v p n x t n r (W tt A GE rth . S th O St rt O 19 h Aph d n r r pr nt d b n r th K r d O hr K n O Ch th O Sn r O 1 n t v p nd v r l dv nt v ( h rd A l nd O C pb ll O Ant p d O nt O 1959 W tt 19 S r b d n l d v n n nd/ r ph t ph E AM E Trox scaber ( nn 17 7 ( 1 p h l l nth n h v p rr n n n d ff r nt t rr tr l h b t t nd d n n (Leptospermum scoparium) r b x d r b nd f r t EMA KS S t d ll l n r b tl r n n rth rn nd th rn r n f r t b lp n r b nd t r v r r l d nd nd p t r l nd SE EC E E E E CES r tt n (195 r n (G v n 195 M n p r n ll p rt nt (19 7 C rl n (1991b S h ltz (19 19 th l rv f d n n pl nt r t ( G v n 195 -3 --3 - h ph d n r nl d n f d r nd n r l M tth (1973 E r n rr tt (1997 E r n v n r h nd d n t n r nf n d® t nd W ll (199 Endröd (197 G v n (195 195 19 h r nd r v r fl dpl n (G v n 1955 h n- 19 19 G v n (195 (19 1 h rd d r b n r pr d n ntl n f r t l f (1959 S h ltz (199 St bn d n (1995 l tt r nd pr b bl f d n d n f b t b rd r pt l W tt (1971 1979 19 nd l r n l (E b r n M tth 1973
A GE rth ® S th ® St rt ® S r E A E I O MIA K r d . hr K n ® Ch th ® Sn r ® r r n pt f th r h v f ll d r n A l nd O C pb ll O Ant p d O nt O t n (1995 h d f n t n Cr n (19 r n l n th S rt d nd ll d n l d d b l l n h r n l nd n l d f r E AM ES A O II AE - Acrossidius tasmaniae ( p 1 7 ( 1 5 Phycochus graniceps r n p rf l S rt d pr t d rrh d nd El t r d h l n t r l t n h p n th p r- 1 ( 1 ; SCA A AEI AE - Onthophagus granulatus f l r t ll n t ll nd r t d ( r n t n h n 1 5 ( 1 7 ; ME O O I AE - Odontria giveni W tt 19 ( 1 Costelytra zealandica 1995 A l d t n l f th r p r ntl (Wh t 1 ( 1 9 Prodontria lewisi r n 19 nd t d b r n et al. (1995 r n r tt n (1991 199 x l d d S rt d ( 15 ; Y AS I AE - Heteronychus arator ( br 1775 ( 151 Pericoptus truncatus br 1775 E n t d nd Cl b d fr El t r f r nd ( 15 d f n d th r p b th f ll n j r h r t r h t r n l f l th l n -l v d l rv nd h rt- EMA KS S t d ll l n r b b tl l v d d lt ll p n d ff r nt h b t t ; rf - d n b tl h f r b tl t v d lt ft n th b d tr l n d nd th pl x pr - th r nt rl n d v r n t b n h l d f n t ; nt nn r r l KEY O SU AMI IES O SCA A AEI AE l bb d ft n rr t ; t x ll x v t d t 1 d th nt r r r n r r l r l r r v f r r r n t d ll ; l p l r n nt n th nth h h xt nd f rth r l t r ll th n ; x tr n l bl ; nt nn th r 9 S p rf l SCI OI EA nt ... 2 C pr thr f l th p t n h n n - d th nt r r r n n t b v ; l p l r n h h th h d tr n l h p n th nd f t n t p r t d fr nth b pr n n d r n t n th pr x r t t rn n th r t n p t n n th r d ; x tr n v r r l htl bl - ( r n r tt n 1991 199 q.v. f r l rv l h r- ; nt nn ft n 1 - nt d ... 3 t r t (1 M t t b h th p l p r ; x bl nd ppr x t d; l tr ll n l n nt r bd n; t ll pr nt; b d ll b- [1 ] l SCI I AE p r ll l ( 1 5 1 APHODIINAE (=CYPHONIDAE =DASCYLLIDAE =DISCYLLIDAE =HELODIDAE) - M t t b h th 1 p l p r; x d t ll 153 15 p r t d; l tr h rt n d l v n p d x- n th 1 5-1 r l f r l 5-5-5 p d; t ll n t pp r nt; b d t t v l r r nd d ( 1 7 SCARABAEINAE IAG OSIS d br n t bl n rr l v l t n rl r nd n tl n d n rr th h rp n l 3(1 d nd pr n t n l pl ; l br l r - r d h h r t n t th pr x h n h d t d xp d; nd bl n l d fr b v f ll d fl x d ( 1 ; lI b t pr n nt ( 1 -15 MELOLONTHINAE Ant nn f l f r r n t h rt nd tr n v r - d nd pr n t n l ll th h rn ft n n l n h d v r d b f n r b nt t b r l r pl x l v t n ; l br br n p b n El tr ft n th r d nd th - n l d; nd bl ll p rtl v bl fr pl t p pl r r x l r nd pr j t n th b v ( 151 15 DYNASTINAE pr t rn l r n r d d; t x l pl t ft n pr nt; th t r l nt l b d b n th SE EC E E E E CES n (19 rr tt (19 rr tt C pb ll (19 r n (19 7 SY O SIS S rt d r ll r pr nt d n th C rl n (1991b r (1993 E b r n l nd f n nd n th th rn t p r t r n n
-31- general. Watt (1982a) recorded 121 nominal native spe- RANGE. North ®, South ®, Stewart O cies. Some 125 species are housed in NZAC, and are Kermadecs O Three Kings O Chathams O Snares O placed in 11 genera. Scirtids are poorly known in New Aucklands O Campbell O Antipodes O Bounties O Zealand; ` Cyphon' particularly is a widespread genus in need of revision. EXAMPLE. Eucinetus stewarti ( r n 1 1 ( ί Adults may be quite common on vegetation near water, 155). and on flowers, and larvae of some species are associated with water in the base of epiphytic plants, e.g., Collo- REMARKS. Suggested colloquial name: plate-thigh spermum hastatum, particularly in wet forest areas. Larvae beetles. of Veronatus and its allies live in humus-rich soil. In general, scirtid larvae are considered to be filter-feeding detritivores (Beier 1952). SELECTED REFERENCES. t (1977a, b), Wheeler & Collecting: beating shrubs, sweeping vegetation in wet Hoebeke (1984). places, sifting organic matter near water, intercept traps, and light traps. [1 ] l C AM I AE RANGE. North ®, South ®, Stewart ® Fig. 156 Kermadecs O Three Kings ®, Chathams ®, Snares O Length 0.8-1.4 mm Tarsal formula 4-4-4 Aucklands O Campbell O Antipodes O Bounties O DIAGNOSIS. d n t l b l r tr n l l with sparse and indistinct pubescence; n p r EXAMPLES. Amplectopus pallicornis Broun, 1886 (Fig. p bl f p rt ll r ll n th lv nt b ll d 153), Veronatus tricostellus White, 1846 (Fig. 154). tr n l d fl x d l r br d thr - rt r f x pr n t l dth Clypeus pronounced. E REMARKS. Sharp (1878) divided the New Zealand pl r d v d d b n l nth into dorsal and scirtids into two groups, those with a distinct antennal ventral halves (genus Clambus). Ant nn 1 - nt d fossa and those without a fossa or with a weak fossa. th - nt d l b S t ll tr n l r l r Suggested colloquial name: marsh beetles. r x pr j t n th r v t p n; t x th xp nd d pl t n l n v ntr t 1 nd f ld d SELECTED REFERENCES. Hannappel & Paulus l Legs short and slender. (1992), LeSage (1991a). SYNOPSIS. Watt (1982a) reported 5 native and 2 adventive nominal species of clambid from New Zealand. [17] l EUCI E I AE Endrödy-Younga (1990) revised the New Zealand clam- Fig. 155 bids, recording 11 species grouped in 2 genera. Clambids may be found in leaf litter, lawn clippings, Length 3.0-3.6 mm Tarsal formula 5-5-5 garden prunings, and compost. Some species are attracted DIAGNOSIS. d tr l n d ll pt l uniformly to light, and can be collected using a UV light trap. brownish to black, th f n d b nt p b n Lawrence & Britton (1991, 1994) reported adults feeding d d fl x d n l d fr b v nd r t n on fungal spores. Kuschel (1990) collected species of n t pr x Ant nn f l f r Eyes small but Sphaerothorax in the Auckland area on tree stumps and in prominent. r th r x r d d El tr t p r n p t- loose moss mats of Polytrichaceae, in organic litter, in a r rl ft n th f n r - tr t n nd l n - heap of Eucalyptus branches and cut bamboo in a sheep t d n l pr d l n n th r d f t r paddock, in a litter sample taken in Pittosporum and r x pr j t n th r v t p n; t x Melicytus scrub, and at the foot of a high coastal cliff. xp nd d th bl t x l pl t nl r d nd p rtl n l n 1 t v ntr t Legs short; tibiae and tarsi RANGE. North ®, South ®, Stewart ® of middle and hind legs bearing combs of dark spines. Kermadecs O Three Kings ® Chathams O Snares O Ad lt r p bl f j p n using their modified hind Aucklands O Campbell O Antipodes O Bounties O legs. EXAMPLE. Clambus domesticus Broun, 1886 (Fig. 156). SYNOPSIS. Eucinetus stewarti, a native species, is the only New Zealand eucinetid, and little is known concern- ing its status and habits. Northern temperate species are REMARKS. Suggested colloquial name: clam beetles. known to feed on fruiting bodies of basidiomycete fungi or on slime mould spores (Wheeler & Hoebeke 1984). SELECTED REFERENCES. Endrödy-Younga (1990).
-3 - Superfamily BUPRESTOIDEA SELECTED REFERENCES. Bellamy & Williams The phylogenetic affiliations of Buprestidae are not (1985), Clark (1938), Dumbleton (1932), Milligan (1974, resolved. The group is currently placed in a monotypic 1985), Morgan (1966), Stuart (1955). superfamily (Lawrence & Britton 1991, 1994). Lawrence (1988) suggested that Buprestidae could be the sister- group of the Byrrhoidea, or might be derived from a Superfamily BYRRHOIDEA dascilloid ancestor. The superfamily concept used here is after Lawrence & Britton (1991). Almost all species of Byrrhoidea are aquatic, semiaquatic, or associated with foliage beside [19] Family BUPRESTIDAE streams. They have 5-5-5-segmented tarsi with the en- larged last segment bearing pronounced claws (except for Fig. 157 Heteroceridae and some Byrrhidae, which have 4-4-4 Length 1.7-10 mm Tarsal formula 5-5-5 segmented tarsi). Adults have a sophisticated type of DIAGNOSIS. Body bullet-shaped, subparallel, pointed wing-folding mechanism, and larvae usually have a single at elytral tip, metallic green or black, often with bright pretarsal seta. yellow spots, more or less glossy, heavily sclerotised, Adults of Byrrhidae, Dryopidae, Elmidae, Limnichidae, appearing glabrous or with scattered hairs. Head strongly and Ptilodactylidae have the following characteristics: deflexed, deeply inserted into prothorax. Prothorax head more or less concealed by pronotum from above; tarsi closely applied to elytra. Antennae moderately long, 5-5-5-segmented, with last segment enlarged; legs re- threadlike to slightly serrate. Mesosternum with a large tractable into body cavities; prosternum with a process cavity for reception of prosternal process; metasternum fitting into a cavity of mesosternum (Fig. 52, 53); body with a curved transverse suture. First 2 abdominal convex dorsally; antennae slender, sometimes weakly or sternites strongly connate, the suture between them distinctly clubbed. The family Heteroceridae is the most vaguely defined. Procoxae globose, their cavities open. divergent in this group, and distinguished by prognathous Tarsi with segments 1-4 lobed below. head, short, thick, sawtoothed antennae, flat, spinose tibiae, and 4-4-4-segmented tarsi, with the last segment SYNOPSIS. Watt (1982a) recorded a total of 4 nominal not pronounced. The family Chelonariidae is also quite species of buprestid, whereas Dumbleton (1932) and distinct, but probably close to the main byrrhid lineage. Clark (1938) had indicated 2 native species and 1 adven- tive. The native species belong to 2 genera of BUPRESTINAE. Adults and larvae of Nascioides enysi are found in [20] Family BYRRHIDAE Nothofagus forests, and are restricted to beech species Fig. 158 (Hudson 1934, Dumbleton 1932, Milligan 1974). In Length 1.8-8 mm Tarsal formula 4-4-4 or 5-5-5 contradiction to earlier beliefs, this species is a symptom rather than the cause of beech mortality. Adults and larvae DIAGNOSIS. Body black, brown, or greyish, sometimes of Maoraxia eremita occur in beech forests, but are not metallic green, broadly oval, short, stout, and usually restricted to beech species. The larvae may be found in thin strongly convex dorsally; dorsal surface usually glabrous dead branches of Nothofagus species (Milligan 1974), and and strongly glossy, rarely dull and pubescent with patches also occur in elm trees, while adults have been collected by of paler hair and with short bristles. Head directed down- beating scrub in summer (Hudson 1934, Dumbleton 1932, wards, more or less concealed from above. Antennae Morgan 1966). 10- or 11-segmented, usually weakly clubbed, with A single flying specimen of Buprestis aurulenta segments 3-7 gradually enlarged. Elytra often pointed collected in Remuera, Auckland, represents a species posteriorly, often with distinct striae. Legs often retract- possibly introduced from the north-west coast of the able into cavities of mesosternum, and prosternum with U.S.A. The larvae bore in old wood of most of the a broad process fitting into cavity of mesosternum (Fig. Californian pines, and are particularly attracted to fire 52, 53). Procoxae transverse, with cavities open. Tarsi scars of exposed pitchy wood (Clark 1938). with segment 3 bearing an anteriorly directed mem- branous appendage below. RANGE. North ®, South ®, Stewart O Kermadecs O, Three Kings O, Chathams O, Snares O SYNOPSIS. Watt (1982a) recorded 49 nominal native Aucklands O, Campbell O, Antipodes O, Bounties O species of byrrhid from New Zealand and over 79, many undescribed, in NZAC. Broun (1910b) recognised 8 gen- EXAMPLE. U ES I AE - Nascioides enysi (Sharp, era (including Nosodendron, now in the family Nosoden- 1877) (Fig. 157). dridae [33]); today there are 8 genera in two subfamilies - Y I AE and SY CA Y I AE Aspects of the higher REMARKS. Suggested colloquial name: jewel beetles. classification of Byrrhidae are discussed by Watt (1971).
-33- Adults have been collected from under stones and on AucklandsAucklands O, Campbell O, Antipodes O, Bounties O C l daisies in the mountains, and by beating moss- covered trees or shrubs; larvae have been found on moss EXAMPLE. rn d r t Broun, 1880 (Fig. 159). and feeding on it (Broun 1910b, Hudson 1934). Kuschel (1990) recorded 2 species at Lynfield, Auckland, collected REMARKS. Suggested colloquial name: hairy water on logs and on fairly damp ground and stream beds with beetles. moss and liverworts. In general adults and larvae occur on mosses or liverworts, and some are found among the roots SELECTED REFERENCES. Brown (1981b, 1991a). of higher plants (Lawrence & Britton 1991, 1994). They may be collected on moss, on the ground, or on tree trunks at night, as they appear to be nocturnal feeders. [22] Family ELMIDAE Fig. 160 RANGE. North ®, South ®, Stewart ® Kermadecs O, Three Kings ®, Chathams O, Snares ® Length 2.8-5.0 mm Tarsal formula 5-5-5 Aucklands ®, Campbell ®, Antipodes O, Bounties O DIAGNOSIS. Body narrowly oval, slightly pointed posteriorly, black, greyish-black, or brownish-black, EXAMPLE. Y I AE - h r h tt n Pascoe, sometimes slightly iridescent, moderately glossy or dull 1875 (Fig. 158). (when glossy often with a metallic sheen), with fine and sparse pubescence, usually of a velvety appearance. REMARKS. The New Zealand fauna is diverse, but is Head small. Antennae moderately elongate and poorly understood at the generic and specific level, and the slender, sometimes appearing slightly bead-like. Eyes group requires revision. prominent and protruding. Pronotum trapezoidal, Suggested colloquial name: moss beetles. with sharp basal angles, narrower than elytral base. Elytra with rows of punctures. Procoxae transverse, SELECTED REFERENCES. Broun (1910b), Gressitt & their cavities open behind. Legs Iarge, stout. Tarsal Samuelson (1964a), Watt (1971). segments simple, the last segment enlarged and with moderately large claws.
[21] Family DRYOPIDAE SYNOPSIS. Watt (1982a) recorded 7 nominal native Fig. 159 species of elmid, but some 16 species are now represented in NZAC, many of them undescribed. All New Zealand Length 3-5 mm Tarsal formula 5-5-5 elmids belong to a single genus of subfamily A AI AE DIAGNOSIS. Body broadly oval, usually more or less Elmids are aquatic or semiaquatic, but none can swim. cylindrical, bluntly triangular posteriorly, brown to Hudson (1934) reported adults of d r p in river black, glossy, covered with long, unruly, protruding beds running over stones at the water's edge, sometimes hairs. Antennae short, with a comb-like asymmetrical taking wing with great rapidity. In general the adults may club of 7 segments (Fig. 29). Pronotum with an be found on stones in river and stream margins, and the impressed longitudinal groove on either side. Base of larvae live under stones near the water. Few larvae have elytra with several impressions. Prosternal process been associated with adults, and careful rearing is needed broadly and deeply received into mesosternum. to provide information for systematic studies. Procoxae transverse, their cavities open. Legs large, thick. Last tarsal segment enlarged and often as long as RANGE. North ®, South ®, Stewart ® 3 or all remaining segments, with large claws. Kermadecs O, Three Kings O, Chathams O, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O SYNOPSIS. Watt (1982a) recorded 4 native species of dryopid in New Zealand, in 2 genera, whereas Hudson EXAMPLE. A AI AE - d r p Broun, 1880 (Fig. (1934) had listed only 2 native species. 160). Adults and larvae of rn d r t may be found in leaf litter, commonly in litter of hard beech ( th f REMARKS. Our elmid fauna is relatively diverse but tr n t forests. Adults of l n l occur under logs poorly known, and there are probably many undescribed in very wet places, and those of r t in river beds, species. The southerrn Larainae are under revision by Dr running over stones at the water's edge in hot sunshine Manfred Jäch (Naturhistorisches Museum, Vienna). (Hudson 1934). Suggested colloquial name: riffle beetles.
RANGE. North ®, South ®, Stewart O SELECTED REFERENCES. Brown (1981 a, 1991 c), Kermadecs O, Three Kings O, Chathams O, Snares O Spangler & Brown (1981).
-3 --3 - [ 3] l IM IC I AE SYNOPSIS. Heterocerids are represented in New Zea- Fig. 161 land by a single native species in the type genus. The type specimen is from Wedderburn in Central Otago, but the Length 1-2.5 mm Tarsal formula 4-4-4 (Hyphalinae), species has also been found at Whatipu and Bethels Beach 5-5-5 (Limnichinae) near Auckland, Ballantrae near Palmerston North, Hau- DIAGNOSIS. d n t t ll v l nv x moana near Hastings, and in Northland. d r t l l black, with short yellowish or greyish All known species live in tunnels in moist sand or mud, p b n ft n f r n v p tt rn n pr n t and feed on algae and organic debris. Our species may be nd l tr r nt l p l t r pr nt; clypeus dis- found in mud alongside streams and rivers, where adults tinct. Ant nn l nd r ft n th 3- nt d l b; may be collected by `shore washing' with water and col- b l t nt nl r d; antennal insertion on lecting floating debris by net. Occasionally large numbers base of frons. r n t l b n t S t ll l r of adults have been caught in light traps near rivers. r x tr n v r th r v t p n behind. Meso- sternum long and wide, well extended into procoxae; RANGE. North ®, South ®, Stewart O t t rn th d t n t tr n v r t r As in Kermadecs O Three Kings O Chathams O Snares O byrrhids, l b f ld d nt v ntr l v t Aucklands O Campbell O Antipodes O Bounties O
SYNOPSIS. Watt (1982a) reported 7 nominal native EXAMPLE. E E OCE I AE - Heterocerus novaeselan- species of limnichid and 1 adventive. The 2 limnichid diae Charpentier, 1968 (Fig. 162). genera in New Zealand belong in subfamilies Y A I AE (5 species) and IM IC I AE (3 species, with possibly a REMARKS. There are related species in Australia. fourth undescribed). Suggested colloquial name: mud beetles. Species of Hyphalus are known from the intertidal zone on barnacle-encrusted rocks and beach rocks. Species of SELECTED REFERENCES. Charpentier (1968). Limnichus occur under rocks at the margin of streams and rivers, and in moist litter in forested areas (e.g., Notho- fagus forest). [ 5] l I O AC Y I AE Fig. 163
RANGE. North ®, South ®, Stewart O Length 5-8 mm Tarsal formula 5-5-5 Kermadecs O Three Kings O Chathams O Snares O Aucklands O Campbell O Antipodes O Bounties O DIAGNOSIS. d bl n t l n t pale brown sometimes with a reddish tinge to dark brown, moderately densely pubescent. d ll tr n l d fl x d nd EXAMPLE. LIMNICHINAE - Limnichus n r p Broun, n l d fr b v ; frontal suture distinct. Ant nn 1 (Fig. 1 1 11- nt d f l f r r rr t d r t l l n (Fig. 20). E l r nd pr tr d n r n t ll REMARKS. Very little is known about New Zealand's tr n v r th ll d v l p d b l l n Limnichidae. h n n t n f b th lI t th r Suggested colloquial name: lake beetles. r n l t n ; l t r l r n h rp nd n pl t nt r rl Elytra striate. S t ll h rt- h p d not- SELECTED REFERENCES. Britton (1973, 1977), ched anteriorly. r x n l tr n v r n rr l Brown (1991b). p r t d th r v t p n b h nd r pl ; t r r ll
[ ] l E E OCE I AE SYNOPSIS. Watt (1982a) reported 2 nominal native Fig. 162 species of ptilodactylid and a total of some 8 species,
Length 3.8-4 mm Tarsal formula 4-4-4 several undescribed, represented in NZAC. As conceived here, the family includes 1 described species of Byrro- DIAGNOSIS. d r b t l n t densely pubescent, cryptus (subfamily A C Y A SI AE with 4-7 awaiting th p b n f r n v r t d p tt rn d description, and possibly another undescribed genus pr n th with strong mandibles and with large (Kuschel 1990). Following the classification proposed by labrum and clypeus. Ant nn h rt nd th th n Lawrence & Newton (1995), Brounia—hitherto placed in l n t pp r ntl 7- nt d l b Pronotum Ptilodactylidae—is transferred to Chelonariidae [26]. strongly transverse. El tr l r l bp r ll l r x Adults are predacious, and occur on vegetation near tr n v r th r v t p n behind. b fl tt n d water in wooded areas. Kuschel (1990) reported adults of p n Byrrocryptus collected by Malaise trap near a stream,
—35 — underneath overhanging stream banks, and amongst fern RANGE. North ®, South O, Stewart O and moss on a steep, seeping slope. Specimens of B. Kermadecs O, Three Kings O, Chathams O, Snares O r h rt have been collected by beating vegetation near Aucklands O, Campbell O, Antipodes O, Bounties O margins of streams. Ptilodactylid larvae from other regions occur in wet habitats, and may be riparian or EXAMPLE. r n th r Sharp, 1878 (Fig. 164). aquatic; they feed on rotting vegetation, including leaves, roots, or dead wood (Lawrence 1991, LeSage & Harper REMARKS. According to Lawrence & Newton (1995) 1976). th r resembles some members of the Callirhipidae more than it does species of Ch l n r or d - RANGE. North ®, South ®, Stewart O h l n r (Chelonariidae), but it shares the serrate Kermadecs O, Three Kings O, Chathams O, Snares O tibiae, tarsal structures, wing venation, and type of male Aucklands O, Campbell O, Antipodes O, Bounties O genitalia typical of chelonariids. Suggested colloquial name: none. EXAMPLE. ΑΝCΗΥΤΑ SIΝΑE — Byrrocryptus urquharti Broun, 1 93 (Fig. 1 3 SELECTED REFERENCES. Lawrence & Newton (1995). REMARKS. The New Zealand fauna of Ptilodactylidae is probably richer than is apparent from its representation in collections. The subfamily Anchytarsinae is almost Superfamily ELATEROIDEA certainly paraphyletic, as suggested by Lawrence & The superfamily definition used here is after Lawrence & Newton (1995). rr r pt which occurs also in Britton (1991, 1994) and Lawrence & Newton (1995). Australia, was placed in this subfamily on the basis of a Adults of all elateroids lack a mandibular mola and a larval synapomorphy—longitudinal division of expanded transverse metasternal suture, and have an unsophisticated submentum into three parts—with four other genera wing-folding mechanism; larvae have a single stemma on unknown in New Zealand (Lawrence & Newton 1995). either side of the head and a feeding mechanism adapted The entire group is much in need of revision. for liquid diets (Lawrence & Britton 1991, 1994). These Suggested colloquial name: comb-waist beetles. plant-feeders usually have the rear of the prosternum elongated into a process that fits into the mesosternum SELECTED REFERENCES. Stribling (1986). (Fig. 54). Phylogeny is discussed by Muona (1996).
[26] Family CHELONARIIDAE [27] Family EUCNEMIDAE Fig. 164a,b [assisted by J. Muona] Length 5-6 mm Tarsal formula 5-5-5 Fig. 165
DIAGNOSIS. Body narrowly oval, partly subparallel, Length 4-10 mm Tarsal formula 5-5-5 moderately convex, in lateral view with an anterodorsal pronotal hump, black. Head small, strongly deflexed, DIAGNOSIS. Very similar to Elateridae [28], but dif- concealed from above; frontal suture distinct. Anten- fering in having the labrum membranous and con- nae 11-segmented, with basal 2 segments small and cealed beneath the clypeus. Body brown to black, segments 4-11 pectinate, asymmetrical (Fig. 22). Eyes narrowly elongate, subparallel, pubescent, broadest at large and protruding. Pronotum small, transverse, pronotum and tapering posteriorly. Front edge of pro- with a welI developed basal locking mechanism con- sternum straight across and not lobed. Head deflexed. sisting of a comb with small teeth; lateral carinae sharp Antennae 11-segmented, filiform or serrate. Pronotum and complete. Elytra with faint indications of striae. with hind angles acutely produced posteriorly. Elytra Scutellum small, oval. Procoxae transverse, their cav- striate. Clicking device present (consisting of prosternal ities open behind; mesocoxae separated by a distance process fitting into mesosternal cavity), clicking ability shorter than coxal width; metacoxae transverse, in contact well developed. Procoxae globular, their cavities open. basally. Metasternal longitudinal suture present and Tarsomeres simple. First 5 ventrites connate. transverse suture absent. Tarsi simple, with segment 3 bilobed beneath and segment 4 minute. SYNOPSIS. Watt (1982a) reported 20 nominal native species of eucnemid from New Zealand. We estimate 22 SYNOPSIS. There is a single native species of chelon- species, including 2 undescribed, in 7 genera and 2 sub- ariid, r n th r in New Zealand. We have families – MELASINAE and MACRAULACINAE. Apart from the examined only 1 specimen, collected in a Malaise trap in scattered original descriptions, very little information is cattle-browsed forest at Hicks Bay, north of Te Araroa. available on the group.
-3 - Adults may be found on forest growth, in leaf litter or well separated, the Iast 2 connected by a membranous wood mould. Kuschel (1991a) reported 2 eucnemid suture. species collected under bark of Pinus radiata logs lying in the bush and in decayed wood in the Auckland area. Hud- SYNOPSIS. Watt (1982a) reported 132 native and 3 son (1934) recorded some species collected by beating adventive species of elaterid in New Zealand, belonging to forest growth. May also be collected using interception 26 genera in 6 subfamilies — AGRYPNINAE, PITYOBIINAE, traps, including Malaise traps, or by sweeping forest DENTICOLLINAE, LISSOMINAE (PROTELATERINAE), ELATERINAE, vegetation. and CARDIOPHORINAE. For subfamily and tribal identification, New Zealand larvae are known from dead wood of see Stibick (1979). Metrosideros (rata). Most Australian larvae also occur in Adults occur on foliage of trees and shrubs, in rotting dead wood (Lawrence & Britton 1991, 1994), and logs, under bark of dead trees, and on the ground, and may Nearctic larvae of the group are known as cross-wood be collected by sweeping vegetation, collecting at light, or borers because they cut characteristic mines across the netting in flight in the daytime. Kuschel (1990) collected grain of wood (White 1983). adults at Lynfield, Auckland from vegetation by beating, from Acacia and Eucalyptus, in bush litter and on low- RANGE. North ®, South ®, Stewart ® growing plants at night, at light, in paddocks, in Malaise Kermadecs O, Three Kings O, Chathams ®, Snares O traps, in leaf litter and prunings, under bark, from sifted Aucklands O, Campbell O, Antipodes O, Bounties O decaying wood, and reared from abandoned boards of Pinus radiata. EXAMPLE. MELASINAE - Neocharis simplex Sharp, According to Lawrence & Britton (1991, 1994), in 1877 (Fig. 165). general there are three types of elaterid larvae: (a) sap- rophagous — feeding on rotten wood; (b) phytophagous — REMARKS. In early classifications the eucnemids were feeding mainly on roots of plants; and (c) predacious. treated as a subfamily of Elateridae [28], but Broun treated Many phytophagous larvae, known as wireworms, are them as a distinct family. Their phylogeny, classification, agricultural pests feeding on roots of cereals and other biogeography, and biology (including New Zealand gen- plants. For example, Ctenicera larvae are soil-inhabiting era) were recently reviewed by Muona (1991, 1993). wireworms, some f them damaging root vegetables such Suggested colloquial name: false click beetles. as carrots. Thoramus larvae live in dead wood, including standing dead trunks of Pinus radiata, where they prey on SELECTED REFERENCES. Muona (1991, 1993), Sharp larvae of Cerambycidae. Most larvae are probably facul- (1877b). tative predators on soil-inhabiting arthropods.
RANGE. North ®, South ®, Stewart [28] Family ELATERIDAE Kermadecs ®, Three Kings ®, Chathams ®, Snares O [assisted by J. Muona] Aucklands O, Campbell O, Antipodes O, Bounties O Fig. 166, 167 Length 3-30 mm Tarsal formula 5-5-5 EXAMPLES. AGRYPNINAE - Thoramus wakefieldi Sharp, 1877 (Fig. 166); DENTICOLLINAE - Amychus candezei DIAGNOSIS. Body elongate, narrow, cylindrical to Pascoe, 1876 (Fig. 167). slightly flattened, usually brown or black, rarely spotted, finely pubescent or nearly glabrous, often glossy, with REMARKS. This group is in need of taxonomic revision, medial part usually subparallel and both ends rounded. as there is no comprehensive treatment available. The Head with labrum sclerotised, visible and free, and general classification of Elateridae is in a state of flux, and clypeus not distinct. Antennae 11-segmented, located the subfamilies are variously combined by different near but not between eyes, usually sawtoothed or comb- authors (Lawrence & Newton 1995). like, never clubbed. Prothorax loosely joining mesothorax Suggested colloquial name: click beetles (some larvae — and capable of rapid movement on basal joint, which wireworms). produces a jumping movement. Pronotum with hind angles acutely produced posteriorly. Click mechanism SELECTED REFERENCES. Calder (1976, 1984, 1996), always present (Fig. 54)—the most distinctive feature of Gaedike (1969), Sharp (1877b), Stibick (1979). the family, consisting of prosternal process and mesoster- nal cavity. Elytra usually subparallel for most of length, tapering posteriorly. Procoxae globose, their cavity open [29] Family LYCIDAE behind Tarsi simple, sometimes with setal brushes or Fig. 168 membranous appendages. Prosternum truncate or lobed in front. Abdomen with 5 ventrites, rarely 6, all usually Length 9-10 mm Tarsal formula 5-5-5
—37— DIAGNOSIS. Body black and orange, soft, flattened, distinctly shortened, exposing wings and many tergites, loosely formed. Head small, partially concealed from often with inconspicuous ridges. Procoxae projecting, above. Antennal bases in proximity; antennae thick, their cavities open. Legs long and slender. Tarsi with strongly serrate, moderately long (Fig. 21). Eyes pro- segment 4 bilobed. truding. Pronotum narrower at base than elytra, with several ridges dividing disc into irregular depressions; SYNOPSIS. Watt (1982a) recorded 40 native and 1 base slightly sinuate. Elytra leathery, narrowly elongate, adventive species of cantharid in New Zealand, belonging subparallel, expanded beyond body margins, with dis- to 3 genera in 2 subfamilies — YSMO OCE I AE and tinct round humeri and longitudinal ridges, between MA I I AE By far the most species (37) are placed in ridges with a double row of large punctures. Procoxae Asilis (Wittmer 1979). A new subfamily classification was projecting, their cavities open behind. Femora and tibiae proposed by Brancucci (1980). slightly flattened. Tarsi with segment 3 and particularly Adults are diurnal and often seen on flowers and foliage segment 4 bilobed. of various trees and shrubs, and occasionally may be found under stones and bark. The majority of species are SYNOPSIS. Watt (1982a) recorded a single species of believed to be predacious on other insects, but some feed lycid from New Zealand, corresponding to Metriorrhyn- on plants, nectar, and pollen (LeSage 1991b). A common chus erraticus Broun (1893a), described from Drury, method for collecting adults is sweeping and beating Pokeno, and Clevedon. This is implicitly a synonym of vegetation. Porrostoma rufipenne, an adventive Australian species, Kuschel (1990) reported 3 cantharid species from reported by Kuschel (1990) from Lynfield, Auckland. Auckland, collected by sweeping grass and weeds, beating Adults fly on warm days, and feed on flower nectar and vegetation, from flowers in a swamp, and in a Malaise trap. p ll n rv b f nd nd r th b τ f l Larvae are primarily nocturnal, preying chiefly on small arthropods living in debris on the ground (LeSage 1991b). A GE rth ® S th ® St rt Ο In New Zealand some larvae have been extracted from leaf Kermadecs O, Three Kings O, Chathams O, Snares O litter using Berlese funnels. Aucklands O, Campbell O, Antipodes O, Bounties O RANGE. North ®, South ®, Stewart O EXAMPLE. Porrostoma rufipenne (Fabricius, 1801) Kermadecs O, Three Kings O, Chathams O, Snares O A l nd O C pb ll O Ant p d O nt Ο (Fig. 168).
EXAMPLE. YSMO OCE I AE - Asilis fulvithorax REMARKS. The bright colours of, e.g., Porrostoma rufi- (Broun, 1880) (Fig. 169). penne may be a warning signal to potential predators, since lycids in general are believed to mimic the colours of REMARKS. Suggested colloquial name: soldier beetles. insects distasteful to predators. Suggested colloquial name: net-winged beetles. SELECTED REFERENCES. Brancucci (1980), LeSage (1991b), Wittmer (1979). SE EC E E E E CES Βο á Β á vá (199
Series BOSTRICHIFORMIA [30] Family CANTHARIDAE We have followed the concept of Lawrence & Britton Fig. 169 (1991, 1994) and Lawrence & Newton (1995) in defining Length 1.9-7 mm Tarsal formula 5-5-5 this group. It is considered to be a paraphyletic assemblage consisting of Polyphaga with strong affiliation to the DIAGNOSIS. Body usually uniformly black or brown, cucujiform lineage, but lacking shared apomorphies with sometimes with pronotum yellowish to orange, soft, the series Cucujiformia. Jacobsoniidae are tentatively narrowly oval to subparallel, slightly convex to flattened, included here because they lack some important syn- moderately densely pubescent, often slightly glossy. Head apomorphies of Cucujiformia (Lawrence & Newton large, deflexed but partially visible from above. Eyes 1995). protruding. Labrum membranous and inconspicuous. Antennae filiform (Fig. 17) or serrate, with insertions welI separated. Pronotum transverse, broadly oval or Superfamily DERODONTOIDEA approximately subquadrate, sometimes with 2 small, Members of this group are distinct in having paired ocelli round basal impressions, and sometimes emarginate. on the frons, an elaborate system of depressions and Elytra narrowly oval, often subparallel for most of usually ridges and canals on head and pronotum, and length, usually loosely covering abdomen but sometimes elytral striae with deep basal pits.
-3 - [31] Family DERODONTIDAE (1991, 1994) and Lawrence & Newton (1995) considered Fig. 170 their placement of Jacobsoniidae in Bostrichoidea to be a tentative arrangement, as at present there is only weak Length 2.0-2.4 mm Tarsal formula 5-5-5 evidence of some similarity between the two groups in DIAGNOSIS. Body small, approximately oval in out- aedeagal characters. line, brown or black, moderately glossy, with incon- spicuous short adherent pubescence. Head with paired ocelli and usually elaborate depressions. Antennae 11- [32] Family JACOBSONIIDAE segmented, the basalmost 2 segments enlarged, and Fig. 171
with a 3-segmented club. Eyes protruding, coarsely Length 0.7-2 mm Tarsal formula 5-5-5 or faceted. Pronotum distinctly narrower than base of elytra, 3-3-3 (appearing 2-2-2) with a series of depressions and usually with ridges dividing disc into several sections. Elytra broadly oval, DIAGNOSIS. Body small to minute, elongate, reddish- with deeply impressed striae each usually having a deep pit brown or brown, glossy, usually punctate, with (unique at base. Procoxae projecting, their cavities usually character) metasternum at least as long as 5 visible closed behind; metacoxae transverse, with well devel- sternites combined, giving the appearance of hind legs oped plates. emerging near the end of the abdomen. Antennae with a 1-3-segmented club. Procoxae projecting, their cav- SYNOPSIS. There are 2 native species of derodontid in ities usually open. New Zealand, both in genus Nothoderodontus of subfam- Of the two distinct groups in our fauna the first, ily LARICOBIINAE (Watt 1982a, Lawrence 1985, Lawrence represented by a single species in genus Saphophagus, has & Newton 1995). the body larger (length 1.9-2 mm) and tarsi 5-5-5-seg- Nothoderodontus gourlayi feeds on hyphae and conidia mented. The second comprises possibly three undes- of sooty moulds (Ascomycetes: Capnodiaceae and related cribed native or introduced species in genus Derolathrus, families) (Lawrence 1985). Specimens including larvae distinct by their minute body size (length 0.7-0.8 mm), have been taken in Nothofagus forest from sooty mould yellowish rust-brown colour, antennae appearing to have a growing on the honeydew excreted by a scale insect. 1-segmented club, head abruptly constricted behind eyes to form short temples, oval pronotum, and narrowly RANGE. North O, South ®, Stewart O oval elytra, tapering distally. Kermadecs O, Three Kings O, Chathams O, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O SYNOPSIS. Watt (1982a) recorded 1 described native species and 2 other species of jacobsoniid housed in EXAMPLE. LARICOBIINAE - Nothoderodontus gourlayi NZAC. These are grouped in 2 genera — Saphophagus, Crowson, 1959 (Fig. 170). with 1 native species known only from the South Island, and Derolathrus, with 1 undescribed species (or possibly REMARKS. Derodontids are considered to be among the 3), collected in Omahuta State Forest, Northland, and on most primitive of Polyphaga (Lawrence & Newton 1995). Codfish Island and Rurima Island in the far south. Dero- Suggested colloquial name: sooty fungus beetles. lathrines are also known from Norfolk Island, Lord Howe Island, Christmas Island, and Australia (Lawrence & SELECTED REFERENCES. Crowson (1959), Lawrence Britton 1991, 1994, Löbl & Burckhardt 1988). Crowson (1985), Lawrence & Hlavac (1979). (1959) considers S. minutus to be one of the most remark- able of New Zealand's endemic insects, possibly a relict coeval with the tuatara, Sphenodon. Superfamily BOSTRICHOIDEA Adults of S. minutus occur under loose bark of dead The concept used here is after Lawrence & Britton (1991, Nothofagus branches. Larvae of S. minutus described by 1994) and Lawrence & Newton (1995). The superfamily Crowson (1959) were found under moist and close-fitting comprises five very diverse families (Anobiidae, Bostri- bark of dead Nothofagus or, in one instance, of pukatea chidae, Dermestidae, Jacobsoniidae, and Nosodendridae), (Laurelia novae-zelandiae). Specimens of Derolathrus all with 5-5-5-segmented tarsi. Lawrence & Newton were found in guano and in a nest of kiore, Rattus exulans. (1982) and Lawrence & Britton (1991, 1994) proposed inclusion of the Dermestidae in Bostrichoidea on the basis RANGE. North ®, South ®, Stewart ® of modified cryptonephridism, similar aedeagal structures, Kermadecs O, Three Kings O, Chathams O, Snares O and lack of a basal mandibular mola in larvae. Ivie (1985) Aucklands O, Campbell O, Antipodes O, Bounties O added Nosodendridae to this superfamily on the grounds of assumed secondary loss of cryptonephridism in both EXAMPLE. Saphophagus minutus Sharp, 1886 (Fig. Dermestidae and Nosodendridae. Lawrence & Britton 171).
-39- REMARKS. The taxonomic position of this small family [34] Family DERMESTIDAE is uncertain (Lawrence & Newton 1995). Fig. 173 Suggested colloquial name: Jacobson's beetles. Length 3-10 mm Tarsal formula 5-5-5 SELECTED REFERENCES. Crowson (1959), Löbl & DIAGNOSIS. Body compact, usually robust, oblong, Burckhard (1988). elliptical, or broadly ovate, usually brown to black, rarely reddish-brown (Trichelodes), sometimes with pale spots, slightly to strongly glossy, clothed with short or long, [33] Family NOSODENDRIDAE erect or decumbent hairs or scales, these often forming Fig. 172 patterns. Head deflexed and more or less concealed from above, usually with a median ocellus. Antennae Length 4-7 mm Tarsal formula 5-5-5 filiform or pectinate, short and clubbed, the club DIAGNOSIS. Body black, ovoid, strongly convex, usually 3-segmented but sometimes 4-8-segmented; glabrous or with sparsely distributed short bristles, antennae often fitting into grooves below either side of moderately to strongly glossy. Head prognathous and pronotum. Eyes moderate-sized to large and protruding. not retractable; clypeus narrow but visible; maxillae and Pronotum transverse, narrower than elytral base, prementum concealed beneath enlarged mentum. Antennae sometimes strongly convex medio-apically, giving a 11-segmented, with a 3-segmented tomentose club. hump-backed appearance laterally, usually with sharp Pronotum strongly transverse (Fig. 118). Scutellum tri- lateral margins (absent in Trichelodini). Procoxae trans- angular, large. Elytra broadly curved laterally, rounded verse, conical or globose and usually projecting, their posteriorly, with punctate or setose striae. Prosternum cavities open; metacoxae usually excavated for with a triangular process fitting into mesosternal reception of femora (Fig. 56). Tarsal segments simple. cavity. Procoxae transverse, their cavities open. Legs capable of folding into ventral cavities. Tarsal segments SYNOPSIS. Watt (1982a) reported 11 native and 6 simple. adventive species of dermestid in New Zealand. They belong in 7 genera representing 4 subfamilies — DER- SYNOPSIS. Watt (1982a) recorded 2 native species of MESTINAE, TRINODINAE, ATTAGENINAE, and MEGATOMINAE. nosodendrid in New Zealand, whereas Hudson (1934) and Species of Anthrenocerus, Anthrenus, Attagenus, Der- Endrödy-Younga (1989) reported 3 species. There are 2 mestes, and Trogoderma especially are economically species represented in NZAC, Nosodendron zealandicum important, and are represented here mostly by adventive and the smaller N. ovatum. cosmopolitan species. According to Crowson (1959) Adults are encountered infrequently in slime fluxes on native New Zealand species represent only 2 of the 6 wounded trees, where they probably feed on bacteria and subfamilies that he recognised, Trinodinae (Trichelodes) products of fermentation. Specimens of N. ovatum have and Anthreninae (Trogoderma). Kuschel (1990) recorded been collected from under rotten branches, from litter, and 8 species from the Auckland area, most with adults taken one from bat guano; two specimens of N. zealandicum from flowers. were taken in a light trap. No habitat data for our larvae are In general adult dermestids usually feed on nectar or available, but Lawrence & Britton (1991, 1994) reported pollen (e.g., Anthrenus, Trogoderma), though some spe- larvae of Australian species living in slime fluxes. cies are known to use the larval food sources or do not take food at all (Booth et al. 1990, Lawrence & Britton 1991, 1994). Larvae are primarily scavengers, feeding on dried RANGE. North ® South O, Stewart O animal remains such as bones, meat or carrion, hides, furs, Kermadecs O, Three Kings ® Chathams O, Snares O leather, wool and woollen products, silk, feathers, dried Aucklands O, Campbell O, Antipodes O, Bounties O insects, spider webbing, nests of birds and mammals, and even wasps' nests (Reesa vespulae — Waller & Watt 1979), EXAMPLE. Nosodendron zealandicum Sharp, 1882 and some are known from stored grain. Some species of (Fig. 172). Trogoderma can subsist partly or entirely on vegetable matter. Larvae of Trogoderma were reported by Crowson REMARKS. Adults of Nosodendridae may be readily (1959) from a number of unspecified New Zealand distinguished from the externally similar Byrrhidae [20] localities, in cavities and under dry bark, mainly of by their prognathous (not retractable) head, distinct cly- Podocarpaceae, and especially rimu (Dacrydium cupres- peus, and enlarged mentum. sinum). Suggested colloquial name: wounded tree beetles. For cosmopolitan pest species of this group see Booth et al. (1990), Bousquet (1990), Harney (1993), and Law- SELECTED REFERENCES. Crowson (1959), Endrödy- rence & Britton (1994). Adventive species in New Younga (1989). Zealand include the carpet beetle, Anthrenus verbasci,
— --— larvae of which can seriously damage untreated woollen The third type (Fig. 176), represented by members of carpets and garments; adults may be found on flowers in the LYCTINAE, is distinguished as follows. Body elongate spring. The Australian carpet beetle, Anthrenocerus aus- but flattened. Head prognathous, visible from above. tralis, has rather similar habits. The hide beetle, Dermestes Antennae with a compact 2-segmented club. Pronotum maculatus, can seriously damage hides and skins. When approximately trapezoidal, broadest apically, with ready to pupate, its larvae bore into wood or other dense distinct lateral carinae. Elytra subparallel, with par- material, often causing damage to structural timbers. allel rows of hairs. Tarsi with basalmost 4 segments equally short. RANGE. North ®, South ®, Stewart O Kermadecs ®, Three Kings O, Chathams ®, Snares O SYNOPSIS. Watt (1982a) reported 1 native, 2 adventive, Aucklands O, Campbell O, Antipodes O, Bounties O and 1 undescribed species of bostrichid (including Lyc- tinae) in New Zealand. Our current estimate indicates 8 EXAMPLE. MEGATOMINAE - Trogoderma maestum species, in 4 genera and 3 subfamilies — DINODERINAE, Broun, 1880 (Fig. 173). LYCTINAE, and EUDERINAE (see Remarks for details). Euderia is represented by a single native species, and the REMARKS. An economically important group with remaining genera by adventive pest species. several serious pests of hides, wool products, museum Adults of Euderia squamosa have been collected by collections, and stored grain products. The dry exuviae beating Nothofagus branches, and larvae have been found and spear-headed hairs of the larvae can cause asthma or in galleries in moist or sappy bark of dead or injured trees. allergies. A detailed description of the bostrichid larva is provided Suggested colloquial name: hide beetles. by Crowson (1961). Elsewhere, bostrichids are known to bore into sapwood and bamboo. SELECTED REFERENCES. Beal (1991), Crowson (1959), Mroczkowski (1968), Peacock (1978, 1993). RANGE. North ®, South ®, Stewart O Kermadecs O, Three Kings O, Chathams O, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O [35] Family BOSTRICHIDAE (incl. LYCTINAE) EXAMPLES. EUDERINAE - Euderia squamosa Broun, Fig. 174-176 1880 (Fig. 174); DINODERINAE - Dinoderus minutus (Fabricius, 1775) (Fig. 175); LYCTINAE — Lyctus brunneus Length 3-7 mm Tarsal formula 5-5-5 (Stephens, 1829) (Fig. 176). DIAGNOSIS. There are three distinct morphological types of bostrichid in our fauna. REMARKS. There are several unreported, probably The first type (Fig. 174), represented by the monotypic recently adventive species of Lyctinae and Bostrichinae in genus Euderia (EUDERINAE), may be characterised as NZAC which might become of economic importance to follows. Body (except pronotum) elongate and slightly the timber industry. Present holdings are: 1 species of flattened; dorsal surface with protuberances and Euderiinae, Euderia squamosa (Fig. 174); 3 species of depressions, and covered with white and brown scales Dinoderinae — Dinoderus minutus (pantropical pest of forming a pattern. Ant nn with a 3-segmented club, bamboo and maize) (Fig. 175), Rhizopertha dominica (a consisting in males of long flabellate branches and in major economic pest in a variety of dried food products), females of shorter asymmetrical segments. Pronotum and one undetermined; and possibly 4 species of Lyctinae narrow, irregularly cylindrical, as broad as head. Eyes —Lyctus brunneus, a cosmopolitan pest in a wide variety of large, protruding. Elytra subparallel, nearly twice as timbers (Fig. 176), L. planicollis, L. linearis, and one of broad at base as pronotum, and with distinct shoulders. uncertain identity. Tarsi with basal segment as long as the following two One of the most interesting bostrichid species in our combined. fauna is Euderia squamosa. According to Lesne (1934, The second type (Fig. 175) is represented by mainly 1938) it represents the most primitive surviving bostrichid introduced species of DINODERINAE (Dinoderus, Rhizo- type. Crowson (1961) considers Euderia to be more pertha), and may be characterised as follows. Body closely related to Bostrichinae than to any other group, and cylindrical, similar to Scolytinae [82]. Head partially or to be regarded as a probable relict of a group from which completely concealed from above by the hood-like the Bostrichinae were derived. Euderia squamosa has also extended pronotum. Antennae with a 3-segmented some anobiid features (Crowson 1961), and was originally asymmetrical club. Pronotum with rasplike teeth at placed by Broun (1880) in the Anobiidae [36]. front. Elytra often obliquely cut posteriorly, and For a key to subfamilies of bostrichid adults, see Crow- usually bearing spines or lateral teeth. son (1961).
—41— S t d ll l n t b r r b tl ( - Ad lt nd l rv b f nd n d d tr l r t n — p d r p t b tl t b r th h p b r nt f rn t r r n t r d pr d t f n l fr t n b d nd dr d r n SE EC E E E E CES n (197 Cr n t r l f n l nd pl nt r n rv r n r ll (19 1 n (193 d nd b r b r r b t l rv f r t n f d n d r f br f n rv f t n n r n r ll f nd n dr r n t r l f n l r pl nt r n [3 ] l A O II AE l f l tt r nd A tr l n p r n n t ( n l I I AE r n n t f b rd nd l n t ( r n 177 17 r tt n 1991 199 S pt n n r d pr d - n th 3-9 r l f r l 5-5-5 p l t n p r n d n r p t f t r d pr d t ll n d nd b b nd n ( th t l Of t l t 3 d t n t bf l n l nd th 199 t 199 rn 1993 On f th dv nt v t n n r r d l d t n h bl b th r p d r-l E r p n pt n n n l nd t n t t nf t pp r n t n n l h v ppr x t d nt nn l t r d f d d n (193 r p rt d d lt f n rt n l l t r l pr n t l r n nd h v p ll t d b b t n f r t r th n t d l p r t d t x In ntr t t An - ( r n t nd fr dr t b r f h (An b b n h v h d d pr th r x ll pl t l r f rn St b p n In l nd An - n l n h d fr b v nd nt t x b p n t t r d t ntr t d tr t r l t b r nd n ll t f rn t r h IAG OSIS d p l br n d r br n r r - v r l p r rd d n th A l nd r b K h l h ll l n t nd l ndr l t v t (199 r ll t d fr llb rd f ld h nd (An b n r t n r p d r-l nd ft n h d d nd f l f n t v nd ntr d d tr nd lt l r d ( t n n l th d th d b nt r hr b nd l f l tt r r t h r r th l th ft n f r n l r p tt rn ( t n n r r l pp r n l br d d - A GE rth O S th O St rt O fl x d nd ll n l d fr b v ( 1 K r d O hr K n O Ch th ® Sn r O Ant nn 9-11- nt d f l f r nd th t A l nd O C pb ll O Ant p d ® nt O d t n t l b ( t n n r ll rr t r p t n t th 3- nt d l b t nt tr l EXAMPLES. A O II AE - Hadrobregmus magnus l n th n d nd xp nd d r t rn r d d ft n (Dumbleton, 1941) (Fig. 177); I I AE - Ptinus p - d pl x v t th v t nt n n n t rn osus Broun, 1880 (Fig. 178). r x l b r pr j t n ; t x ll nt x v t d t r v f r nd th x l EMA KS A rd n t Iv (19 5 th An b d pl t n An b n n t x v t d nd th t x l h ld b n d r d bf l f tr h d [35] pl t n t n n Abd n th ll 5 v ntr t r nf r t n n th p f n p rt n nt n (19 1 th t l (199 t (199 SY O SIS h rld f n f n b d ( n th n f nd rn (1993 r n r tt n 1991 199 pr 1 S t d ll l n b r r b tl f rn t r n r th v r p ; th r h h r l f t n b tl ( t n n — p d r b tl r r r v n A d rn bf l rr n nt pr v d d b r n t n (1995 nd t th SE EC E E E E CES E p ñ l (197 b 1979 bf l f An b d f th rld ( ld r n pt n (197 197 f M ll n (1977 1979 Wh t x l d n t n d pr v d d b Wh t (197 (197 In l nd th r r 3 n t v nd 5 adventive n n l p f A O II AE nd O CA OMI AE ( r - n ll n An b d nd 5 n t v nd adventive p S r CUCU I O MIA f I I AE ( r n ll n t n d (W tt 19 ; th W h v f ll d th n pt f r n r tt n b l n t 5 n r K h l (199 r rd d 9 (1991 199 nd r n t n (1995 n d f n n p f An b d n l d n 1 pt n n fr nf ld th r Ad lt r h r t r d b n r pt - A l nd v p n n r h v b n r rd d n phr d l M lp h n t b l l f f n t n l p r l fr th K r d I l nd th xtr l t l r rd n bd n l nt t x n t x v t b n fr rf l I l nd nd rd I l nd (E p ñ l f t x l pl t r d t n f pr n t l nt 9 1979 t n t t h b n r rd d fr th A l nd nd 1 nd th f r f th l n t l rv h v I l nd Ant p d I l nd C pb ll I l nd nd h n pr t v p l ph n f t r ; ll th l nd Sn r (W tt 19 1971 l n r f d nt n l l
- - S p rf l C E OI EA p n 1 n r nd x pt f r th dv nt v Ch r t r d b d lt th d r f b d ll t r d pr d t p t n br d r t n v r d th pr tr d n br tl h r pr j t n pr x M t p f r t d r pr d l rv nd 5- nt d t r th nt ft n l b d nd nd d lt ( r n r tt n 1991 199 b t pr b bl l rv th n l pr t r l t l f nd b l r l ll ( r t l t d lt f th tr b nt n n r n t n ll t nd ll pr d h b t ( r n h ll r n l f l tt r f n t v f r t nd r r tt n 1991 199 r n t n 1995 rv l b r r n fl r f n r d t ( n r rd ; t ll h v b l nd b l r pr nd p d n l t l t f th f d n p ll n (Cr n 19 h t n th l ( r n t n 1995 n l n n r nt n n l rv p t d t p n A t th f l f Cl r d ( d lt nd l rv rb r l h b t t (Cr n 19 h r n n p pr v d d b Cr n (19 ll pr b bl b t n nd r b r f d d tr rv f n p f p d pt r x (f r rl p r n r pr d t r n th h b t t nd r n [37] l OGOSSI I AE t n th th d lt (In l E I AE C ll t n ft n l f l tt r nd pr n t thr h 179-1 1 rl f nn l b t n f r t f l r p n d lt fr nd r b r K h l (199 ll t d ht p f n th -1 r l f r l 5-5-5 nt n n n A l nd fr v t t n d d d IAG OSIS W h v f ll d th l f t n pr - r nd l tt r nd tr -b d p d b r n t n (1995 G n r l h r t r t nt nn 11- nt d A GE rth ® S th ® St rt th l 3- nt d l b t pp r n K r d O hr K n ® Ch th ® Sn r O tr l; pr x l v t p n r l d; b- A l nd O C pb ll O Ant p d O nt O d n th 5 v bl nt hr n b d f r b r n d n r tr t d f n f ll E AM ES ΕΝΤΟΝΙΙΝΑE - Rentonidium t v ntr Cr n 19 ( 179 ; LOPHOCATER1NAE - Gr n O O E I AE b d d - z d (l n th - r l r r n 1 ( Ι ; TROGOSSITINAE - f n t d l d f r br dl v l bp r ll l Lepidopteryx nigrosparsa (Wh t 1 ( 1 1 d ll v r d th l n h r ; nt nn th tr l v l 3- nt d l b; l tr tr n l EMA KS h h h r l f t n f r t d p n t t b t th t tr n t ttl d An p rt nt t p n l f t n f th r p E O II AE ( 179 ll t n t b tl th t bl h nt b Cr n (19 19 f th (l n th -1 p rf ll l r t l b d b t tr t d pl x th n th p rf l Cl r d r d l d t n h bl b th l f t x l pl t ; Cr n (19 x r n d f l lt d h v n b d v l t n rl ph r l tr n l nv x nd bf l f h h nt n n r t d f r 5 l l br r th h rt nd p r p b n ; l nd n r S b ntl Cr n (197 d v d d l r br n t bl r t p l r dd h br n; th tr t d ( n br d n nt 3 d t n t f l h d br d nd d fl x d; pr n t br dl r nd d d t n h n th lt d nd r t d b d ff r- nt r rl th fr nt n l n t pr d d; t ll n n th r thp rt th l tt r r p h v n d lt d r t l l r nd bl th t d f n t l r p rt nd th l n O OCA E I AE nd OGOSSI I AE ( 1 1 1 th t n p l h nd l rv l nd bl th lI t d - z d b tl (l n th 3-1 d ff r nt l n nd b l nd th x ll r l th p rf ll l r t t n br n d b t r d l p d n l t t h l f t n f r t d nd d t n h bl b 5-5-5- nt d t r nd th r r l t n h p f th h h r t x f Cl r d r d d h r t r ; b d br dl t n rr l v l; l tr ft n b rr n (1971 nd Sl p n (199 r n bp r ll l fl tt n d t d r t l nv x; pp r r tt n (1991 199 r d th 3 tr t d f l f rf th h rt r l n h r br tl r l Cr n (197 nt th n l f l r t d ft n b l r d nd p tt d; l r br n t bl th S t d ll l n d ll b tl (p lt n - p l r p t ( r h ll h t d r t l l t h ld b tl d ll; h d d r t l n rr nd pr n th ; pr - n t ll th nt r rl pr d d fr nt n l KEY O SU AMI IES O OGOSSI I AE OCCU I G I EW EA A SY O SIS r n t n (1995 r n tr - [M d f d fr Sl p n 199 ] t d pr n 9 bf l f h h r r pr nt d n l nd - O O E I AE E O II AE 1 r x l v t xt rn ll l d O OCA E I AE nd OGOSSI I AE O r f n f ( 1 1 OGOSSI I AE
— 3—
—Procoxal cavities externally open Suggested colloquial name: southern cadelle beetles.
2(1) Protibia with 1 spur; antennal club markedly asym- SELECTED REFERENCES. Watt (1979b).
metrical (Fig. 180) .. LOPHOCATERINAE —Protibia with 2 unequal spurs; antennal club sym- metrical 3 [39] Family CLERIDAE Fig. 183, 184 3(2) Very convex and rounded, clambid-like; procoxal Length 5-20 mm Tarsal formula 5-5-5, cavities internally closed (Fig. 179) .. RENTONIINAE —Never very convex and rounded, not clambid-like; appearing 4-4-4 procoxal cavities internally open PROTOPELTINAE DIAGNOSIS. Body elongate, usually subparallel, often subcylindrical, with dorsal surface usually coarsely SELECTED REFERENCES. Crowson (1964, 1966a, punctate. Coloration dark to pale brown, black, violet, 1970), Foster (1991a), Slipinski (1992). yellow, often spotted with yellow or pale brown, rarely multicoloured, often with a metallic tinge. Setation of variable length, short and decumbent or long and erect, [38] Family CHAETOSOMATIDAE uniformly brown or yellow, and often forming a colour Fig. 182 pattern. Head slightly deflexed, usually insignificantly
Length 7-14 mm Tarsal formula 5-5-5 broader than pronotum. Antennae short, sawtoothed or filiform, most often with a 3-segmented club consisting DIAGNOSIS. Body rectangulate, at least 4x longer of elongate and often asym-metrical segments. Eyes large, than broad, subdepressed, coarsely punctate, glossy, often protruding, slightly to strongly emarginate. with exceptionally long and protruding hairs (often Labial palps with terminal segment often enlarged. Pro- longer than width of elytron). Head strongly prognathous, notum usually narrower than elytral base (exception: with pronounced mandibles. Antennae 11-segmented, r b broadest medially and of equal width basally uniform in width, about as long as forebody, the and apically, usually without lateral carinae (with carinae segments bead-shaped. Eyes small but slightly protru- in M t x n Elytra subparallel, pear-shaped or grad- ding. Pronotum subquadrate, in our fauna without setal ually broadening apically (exception: abbreviated in tufts; front angles pointing slightly anteriorly. Elytra pr irregularly and more or less coarsely punctate. subparallel and elongate, with humeral angles acute Procoxae projecting, their cavities open or closed. Legs (Ch t or obtuse (Ch t d and with long- long. Mesotrochanters oblique (Fig. 58). Tarsi usually itudinal rows of punctures. Scutellum oblong. Procoxae with 1 or 2 segments lobed (simple in M t x n basal globose and slightly transverse, their cavities open. segments sometimes reduced. Abdomen with 5 or 6 Legs with distinct trochanters. Tarsal segments simple. ventrites. Pygidium slightly exposed. SYNOPSIS. Watt (1982a) reported 37 native and 2 SYNOPSIS. Chaetosomatids are known to occur in New adventive species of clerid in New Zealand. These rep- Zealand and Madagascar (Ekis & Menier 1980). There are resent some 8 genera in 6 subfamilies – THANOCLERINAE, 2 endemic genera in New Zealand, Ch t with 1 PHYLLOBAENINAE, CLERINAE, ENOPLIINAE, TARSOSTENINAE, described species and 2 undescribed (NZAC) and Ch t - and KORYNETINAE. h t ph is the largest New d with 1 described species. Zealand genus, containing the majority of species. Our Adults and larvae may be found under bark of dead endemic genus M t x n the most divergent, and there is trees, logs, etc., where they feed on other wood-boring some confusion as to its phylogenetic affiliations; it insects. The larva of Ch t r t d inhabits bur- probably represents a primitive lineage, and may be rows in th f and other trees. considered a relict in the New Zealand fauna (Crowson 1964). Kuschel (1990) collected 5 clerid species in Auck- RANGE. North ®, South ®, Stewart O land, including the 2 adventive ones. Kermadecs O, Three Kings O, Chathams ®, Snares O Adults of many New Zealand native species are Aucklands O, Campbell O, Antipodes O, Bounties O commonly found on flowers and leaves in sunny locations, where they prey on other insects. The predominant habitat EXAMPLE. Ch t r t d Westwood, 1851 of larvae is dead wood, and their food consists of larvae of (Fig. 182). woodborers such as Anobiidae, Bostrichidae, Buprestidae, Cerambycidae, or Scolytinae (Crowson 1964). The cos- REMARKS. Their present widely disjunct distribution mopolitan species r b r f ll and r f p occur suggests that chaetosomatids are an old group of Gond- on carrion and stored products of animal origin, and have wana origin. been reported as feeding on a cargo of copra (Watt 1975).
–44– C ll t n n tt n d lt n fl ht p n v t t n EMA KS r t th d lt f ll d r b d p rt l rl h n n fl r nd ft n b r nd f r t p f Phycosecis, Cr n (19 d br f r l rv S t d ll l n b h b tl
® A GE rth ® S th St rt SE EC E E E E CES Cr n (19 W tt K r d ® hr K n ® Ch th ® Sn r O (1975 19 3 A l nd O C pb ll O Ant p d O nt O
EXAMPLES. THANEROCLERINAE - Metaxina rn t Broun, [ 1] l ME Y I AE 19 9 (Fig. 1 3 ; Ε OΡ II ΑE - Phymatophaea violacea ( l ASY I AE (Fabricius, 1775) (Fig. 1 1 EMA KS A n r t d lt nd l rv f n th 5- r l f r l 5-5-5 l nd t x pr v d d b Cr n (19 th IAG OSIS d ft bl n t n rr l l n t t th bf l f d lt Cl r d r p l t n d r ll fl tt n d ll th d b nt h r nd p f n p rt n th et al. (199 ft n th dd t n l tt r d r t t n f r l S t d ll l n h r d b tl d r - l r d n bl r br n r n h r bl ft n th t ll t n ll l ; d r l rf SE EC E E E E CES Cr n (19 t r f f r b d t th h d r lpt r (1991b d ft n d pr d d ll n t r r l r t- r t ll p r bl n z t pr n t br ll l r t d; nt f x ll r p lp n t nl r d [ ] l YCOSECI AE Ant nn ll f l f r th t l b E nt r 1 5 d r t l l r l htl pr tr d n r n t d - n th 5-3 r l f r l 5-5-5 t n tl n rr r th n l tr l b ( l htl br d r th n dth f l tr n ft n br d t t ddl ft n th IAG OSIS d l n t v l nv x v r bl n d pr n El tr nt r l v r n bd n r l r tl d lI bl b t th t f pr n t l htl h rt r xp n bd n l p x bp r ll l nd h d b r n h t h l d pr n th nd r nd d p t r rl r r d ll br d n n t rd p rt ll n l d b r l r nt r r r n f p x r x pr j t n th r v t p n pr n t th n l r pr j t n b t n nd l n nd l nd r r l nt pl ; t r n l b f nt nn Ant nn pp r n 1 - nt d nt b r n l ll th br n th 1- nt d l b ( l b t ll pp r - pp nd Abd n th v ntr t nt d r n t h r h - h p d n tl n th fr nt n l tr n l pr d d nt r rl El tr SY O SIS W tt (19 r p rt d 33 n t v p f v l th p r l d tr b t d th br tl nd l r d n l nd ll n bf l ASY I AE n b nt r x tr n v r th r v t h r r 3 d r b d n r nd 1 nd r b d nd l d b p n r l nt pl M t - n M t p b l n n `Dasytes' r n Arthra- t rn h rt Abd n th 5 v ntr t canthus, th t t f h h n t l r Ad lt r nl h rb v r nd b f nd n SY O SIS h d r n n fr l nd fl r n hr b h r th f d n p ll n rv f C l d n n t nd A tr l ( r n l nd p r n l f l tt r n d d tr -f rn t n 1995 h f l pr nl n l n r h d nd nd r b r (Cr n 19 K h l (199 Phycosecis, th 1 p rr n n l nd nd ll t d v n p n A l nd fr v t t n 3 th r r p rt d fr A tr l ( r n r tt n fl r nd n M l tr p 1991 199 W tt 19 b 19 3 Ad lt nd l rv f P. limbata r n n nd A GE rth ® S th ® St rt b h nd t l d n th f d n ll b h K r d O hr K n ® Ch th ® Sn r O rthr p d nd r fr ntl n n d d f h r b rd A l nd O C pb ll O Ant p d O nt O
A GE rth ® S th ® St rt ®? E AM E ASY I AE - `Dasytes' subcyaneus r n K r d ® hr K n O Ch th ® Sn r O Ι ( 1 A l nd O C pb ll O Ant p d O nt O EMA KS M j r (199 pr p d n f l - r p E AM E Phycosecis limbata ( br 17 1 ( l f t n f M l r d nd r l t d f l b d n 1 5 l d t n l n h h f r r t n r l v t d
— 5— to family rank. The group is badly in need of revision. dripping sap, vegetable debris, fermented fruits, on fungi, Suggested colloquial name: flower beetles. and in stems of dead fronds of tree ferns. Many species are known to feed on sap and fermented juices. Kuschel SELECTED REFERENCES. Crowson (1964), Foster & (1990) recorded 16 species in 8 genera from Auckland, Lawrence (1991), Majer (1994). from fermenting fruits and vegetables, garden litter, lawn clippings, old logs, trunks of puriri ( t x l n with sap, dry carrion, and sooty mould on manuka ( pt - Superfamily CUCUJOIDEA (CLAVICORNIA) p r p r Introduced C rp ph l species The group concept and family arrangement used here is occur in fallen fruits in orchards, and may transmit yeasts similar to that of Pakaluk t l (1994), Lawrence & causing fruit degradation. Britton (1994), and Lawrence & Newton (1995). The family name Lathridiidae is replaced by its older synonym RANGE. North ®, South ®, Stewart Corticariidae. Members of Cucujoidea are characterised Kermadecs ®, Three Kings O, Chathams ®, Snares O by females having a tarsal formula other than 5-5-4, Aucklands O, Campbell O, Antipodes O, Bounties O distinctive male genital configuration, and larvae approx- imately cylindrical, with a pygopod-like 10th abdominal EXAMPLE. NITIDULINAE - l t p d p r ll Broun, segment (Lawrence & Britton 1994). Adults are generally 1893 (Fig. 187). small, and most have clubbed antennae. REMARKS. This family is very poorly known in New Zealand, and is in need of revision. [42] Family NITIDULIDAE Suggested colloquial name: sap beetles. Fig. 187 SELECTED REFERENCES. Dobson (1993), Kirejtshuk Length 2-6 mm Tarsal formula 5-5-5 (1990), Pakaluk t l (1994). DIAGNOSIS. Body narrowly to broadly oval, or narrowly subparallel ( r h p pl moderately con- vex to flattened, glossy to matt (S r n glabrous or [43] Family MONOTOMIDAE with short and sparse pubescence, or with bristles Fig. 188, 189
(S r n smooth or with tubercles (S r n Coloration Length 2-6 mm Tarsal formula 5-5-5 brown to black, or pale brown; elytra and often pro- or 5-5-4 (males) notum with yellowish spots. Head usually abruptly constricted at base of clypeus and behind eyes; fronto- DIAGNOSIS. Body narrowly oval and moderately clypeal suture almost always absent. Antennae with an convex (M n t to elongate, subparallel and sub- abrupt, usually 3-segmented, ball-like club (Fig. 23). cylindrical ( n x nearly glabrous with some vestigial Lacinia usually without a galea. Mandibles often project- pubescence to scarcely and finely pubescent. Head ing. Pronotum transverse, often emarginated apically, abruptly constricted behind eyes, forming distinct with anterior angles often projecting. Elytra usually temples; frontoclypeal suture absent. Antennae ap- short, exposing pygidium and some tergites or entirely pearing 10-segmented, and antennal club appearing covering abdomen (e.g., l t p d rounded apically, 2-segmented with a distinct seam between club seg- truncate, or split with apices sinuate (some Ep r ments (club actually might be 1-segmented with 2 distinct Procoxae transverse, with exposed trochantins, their areas). Eyes coarsely faceted. Pronotum elongate, with cavities open or close behind. Tibiae often expanded and anterior angles more or less produced and lateral carinae spinose. Tarsal segments 1-3 usually lobed, and distinct. Elytra shortened, exposing at least 2 distal segment 4 sometimes reduced. segments (M n t or only the pygidium ( n x with punctate striae or ridges. Procoxae globose and separate, SYNOPSIS. Watt (1982a) recorded 20 native and 8 their cavities closed. Tarsi with the last segment en- adventive species of nitidulid in New Zealand. We larged, as long as remaining segments combined. First recognise 21 native and 11 adventive species (not all ventrite as long as the next 2 combined. necessarily described) in NZAC, assigned to some 12 genera in 4 subfamilies — CARPOPHILINAE, NITIDULINAE, SYNOPSIS. New Zealand has a single native monotomid, CILLAEINAE, and CRYPTARCHINAE. For subfamily classifi- n x r nd and 5 adventive European species of cation see also Kirejtshuk (1986, 1990). The cosmopolitan M n t (Watt 1982a), all in subfamily MONOTOMINAE. genus C rp ph l contains several species of field and The native species, originally found in Peel Forest, stored products pests. Canterbury, has since been recorded from native forests Adults are found in a diverse range of habitats, on both main islands. Adults of M n t have been including forest growth, flowers, wounds in trees with collected mainly in the Nelson and Auckland areas.
—46— Kuschel (1990) recorded 4 species of Monotoma from explanate, with lateral margins complete. Elytra broad Auckland. basally, tapering posteriorly, with sides slightly ex- Adults of Lenax mirandus have been collected from planate. Hind wings absent. Legs long. Tarsal segments under bark of dead trees of Nothofagus fuscus and N. simple. Procoxae broadly separated (Fig. 55) and glob- menziesii, and in dead wood and/or Platypus tunnels in ular, their cavities open. heketara (Olearia rani). Adults of Monotoma spinicollis in the Auckland area were found in mixed compost, lawn SYNOPSIS. Two subfamilies occur in New Zealand, clippings, garden litter, hen-house straw, and sheep each represented by a single genus and species. Adults and shelters in a paddock (Kuschel 1990). larvae of Agapytho foveicollis (AGAPYTHINAE) live in a black sooty mould which grows on the honeydew excreted RANGE. North ®, South ®, Stewart Ο by scale insects on the trunks of beech trees (Nothofagus), Kermadecs O, Three Kings O, Chathams O, Snares O or in moss and leaf litter. Those of Priasilpha obscura Aucklands O, Campbell O, Antipodes O, Bounties O ( IASI I AE occur in leaf litter of beech forests.
EXAMPLES. MO O OMI AE - Lenax mirandus Sharp, RANGE. North ®, South ®, Stewart O 1877 (Fig. 188), Monotoma spinicollis Aubé, 1857 (Fig. Kermadecs O, Three Kings O, Chathams O, Snares O 189). Aucklands O, Campbell O, Antipodes O, Bounties O
REMARKS. Suggested colloquial name: none. EXAMPLES. AGA Y I AE —Α p th f v ll Broun, 1921 (Fig. 190); IASI I AE - Priasilpha b r SELECTED REFERENCES. Kuschel (1979), Pakaluk et Broun, 1886 (Fig. 191). al. (1994). REMARKS. The status of Phloeostichidae is unsettled. Sen Gupta & Crowson (1969b) redefined the concept of [44] Family PHLOEOSTICHIDAE the family, discussed larval characteristics, and provided a (= IASI I AE key to the subfamilies. Fig. 190, 191 Suggested colloquial name: bark row beetles. Length 3-5 mm Tarsal formula 5-5-5 or (males) 5-5-4 KEY TO SUBFAMILIES OF PHLOEOSTICHIDAE KNOWN FROM NEW ZEALAND A diverse assemblage of primitive cucujoids, which may be variously rearranged after adequate revision. Pakaluk et Winged, with metasternum not very short and metacoxae al. (1994) and Lawrence & Newton (1995) listed 6 sub- narrowly separated; protrochanter partly occluded; families of Phloeostichidae in the world fauna. The New head with a weak postocular constriction Zealand fauna belongs to 2 subfamilies, as follows. ... (Fig. 190) .. AGA Y I AE —Apterous, with metasternum very short and metacoxae DIAGNOSIS. AGA Y I AE (Fig. 190): superficially sim- widely separated; protrochanter fully exposed; head ilar to salpingids, but distinguished as follows. Body without a postocular constriction narrow, moderately convex. Head with a distinct neck, ... (Fig. 191) .. IASI I AE nearly as broad as maximum width of pronotum. Clypeus large. Antennae bead-like, with a weakly defined 3- SELECTED REFERENCES. Crowson (1973), Pakaluk segmented club. Eyes protruding. Pronotum slightly et al. (1994), Sen Gupta & Crowson (1969b). narrower than elytra, with lateral carina incomplete anteriorly, broadest at middle and with 2 deep foveae on either side of base. Elytra oval, in outline, the shoulders [45] Family SILVANIDAE slightly produced anteriorly, with 2 oblique depressions Fig. 192, 193 basally. Hind wings present. Scutellum transverse. Pro- Length 2.5-9 mm Tarsal formula 5-5-5, coxae separated, slightly globose and transverse. Legs appearing 4-4-4 long. Tarsal segments simple. IASI I AE (Fig. 191): superficially similar to sil- DIAGNOSIS. Body small to medium-sized, narrowly phids and agyrtids, but distinguished as follows. Body elongate and subparallel, slightly flattened to weakly broadly oval and slightly depressed. Head small, par- convex, moderately glossy to matt, with inconspicuous tially retractable into pronotum, without an apparent short, decumbent pubescence. Colour brown or rust neck, strongly narrower than maximum pronotal brown. Head prognathous, abruptly constricted poster- width. Antennae with a distinct 3-segmented club. Eyes iorly to form temples. Antennae usually 11-segmented, protruding. Pronotum strongly transverse and laterally without (native species, Fig. 192) or with a 3-segmented
—47— l b ( ntr d d p 193 ; n rt n ft n d ff r n fr th t f r n r tt n (199 nd h dd n fr b v b d f fr n E d t n t r n t n (1995 r l f t d r n t ll l n t (tr n v r S lv n d b tl r ft n nf d th C j d / n Ahasverus), nd ft n th pr j t n fr nt n l phl d [ 7] nd r t r rd d nd/ r d nt t r r n l t l t r l r n ; d bf l f th t r p ( W tt 19 C pl t I f t th l n t d n l d pr n El tr b- th ` h ld b h lpf l t p r t th p r ll l br d r th n pr n t nd th d t n t S t d ll l n n n h ld r ll tr t l p n t t r x ll l b n rr l p r t d nd p n r l d r l SE EC E E E E CES l t d (1973 19 nt 3 b l b d ( x pt n r nt n nd nt f v t h (19 1 l et al. (199 h (19 ll ll v n th pr n f p d t tr r nt t n [ ] l CUCU I AE SY O SIS W tt (19 n l d d lv n d bf - 19
l f C j d [ ] nd r p rt d th n b r f n n l n th 15- r l f r l 5-5-5 p n t v nd dv nt v ll n l et r 5-5- ( l al. (199 nd r n t n (1995 r n lv n d nd j d d t n t f l nd f nd 7 IAG OSIS d l n t - v l bp r ll l tr n l n t v nd 5 dv nt v lv n d r pr nt d n AC In fl tt n d l br nd d r t l l d l r l nd lv n d r r pr nt d b bf l l htl br d r th n pr n t ; l br n l d BRONTINAE nd SI A I AE h th n r Ant nn 11- nt d n l f r ; b l nt O r p f r nt n l v nd r b r f d d n t v br dl l n t nd nt ll nd b d h p d tr r t p n tr r n l f l tt r Ad lt nd 3rd n n rr l l n t nd th l n t 7th nd th l rv f Cryptamorpha brevicornis ft n r t th r l htl l n t nd 3 t r n l nt b d h p d nd r l b r f d d tr nd n th b f d d v n th pr n f l b E ll b t f ll n l v f n p l (Rhopalostylis sapida). l htl pr tr d n r n t tr n v r th rr t S lv n d r n n t f d pr r l n d d pl nt d t n t l t r l r n ; fr nt n l l htl pr d d t r l nd f n b t r dv nt v p r t r d nt r rl El tr pl t l v r n bd n b- pr d t p t r n t n th p l t n Ahasverus p r ll l n b l t -th rd th r l v bl tr advena f nd fr ntl n t r d r n n t nd nd th pl t l t r l r n r x l b b n h r t f d n ld nd pl nt d br nd br dl p r t d b pr t rn l pr th v t Oryzaephilus surinamensis n n ll p rt nt p n; x l b br dl p r t nd p n; p t f r l dr d fr t nd l d Ad lt f t x tr n v r p r t d p ll b t n nt t Nausibius clavicornis r ntr d d th A tr l n b ll r n r d t l n th n ll p rt nt lv n d p th et al. (199 t (199 nd SY O SIS C j d r r pr nt d n l nd rn (1993 nl b n nd r b d p f Platisus nd t th r nt n b ll t d b p n p n fr hr K n I l nd f r t r rd d b W tt (19 M t nd r b r b ft n f r t l tt r nd pr n t thr h p n r ll t d n v b r nd br r fr rl f nn l r n p tf ll tr p n n t v b h r nd r b r f n (Kunzea ericoides), p ll l r r f r t r ll t n d t l f lv n d p ld tr n r x d f r t r n nt rr n n A l nd K h l (199 A GE rth O S th O St rt O A GE rth ® S th ® St rt O K r d O hr K n ® Ch th O Sn r O K r d ® hr K n O Ch th O Sn r O A l nd O C pb ll O Ant p d O nt O A l nd O C pb ll O Ant p d O nt O E AM E Platisus p ( 19 EXAMPLES. Β O I AE - Brontopriscus pleuralis EMA KS h n p fr th hr K n (Sharp, 1877) (Fig. 192); SI A I AE - Oryzaephilus th nl r pr nt t v f th f l C j d n surinamensis (Linnaeus, 1758) (Fig. 193). l nd A rd n t W tt (19 r pr nt t v f Platisus pr b bl n rr d n th l nd EMA KS l t d (1973 19 r v d th n r nl nd nd rv v d l l l t nl n th n rth Silvanus nd Oryzaephilus, nd d d b l ll n th r l t v l ldl ff t d hr K n b n p rt nt p f v t h (19 1 pr p d l - xt n t n th nl nd S v r l p f Platisus r f t n f n r n th t l nd f l n n fr t rn A tr l ( r n r tt n 199
— — In the past, members of the Silvanidae [45] and Laemo- EXAMPLE. Microbrontes lineatus (Broun, 1893) (Fig. phloeidae [47] were erroneously treated as cucujids. 195). Suggested colloquial name: flat beetles. REMARKS. Thomas (1995) recognised Microbrontes, SELECTED REFERENCES. Pakaluk et al. (1994), Watt Cryptolestes, and Placonotus—males of which possess (1986). various modifications to the antennal scape—as members of family Laemophloeidae (Cucujidae in a broad sense). Pakaluk et al. (1994), Lawrence & Britton (1994), and [47] Family LAEMOPHLOEIDAE Lawrence & Newton (1995) recognised Cucujidae and Fig. 195 Laemophloeidae as distinct families. Suggested colloquial name: none. Length 1.8-3 mm Tarsal formula 5-5-5 or (males) 5-5-4 SELECTED REFERENCES. Lefkovitch (1958, 1961), Includes all the smaller, flattened cucujoids with submar- Pakaluk et al. (1994). ginal carinae on the pronotum.
DIAGNOSIS. Body narrowly elongate, strongly flat- [48] Family PHALACRIDAE tened, glabrous or with fine and sparse pubescence, Fig. 196 dark brown to yellowish or reddish-brown, moderately to Length 2-3 mm Tarsal formula 5-5-5 or 5-5-4 densely punctate. Head flat, as broad as pronotum or slightly narrower. Labrum concealed. Antennae 11- DIAGNOSIS. Body small, approximately hemispher- segmented, moniliform, without a distinct club; males ical, flat ventrally, variably punctate, at x40 magnification with various modifications of the scape, in Microbrontes appearing glabrous, strongly glossy, red-brown or taking the form of a mesally directed horn or process. dark brown to black, with a metallic sheen; appendages Eyes small. Pronotum with a sublateral carina on either paler and usually reddish-brown. Head short, broad, side; anterior angles often acute. Elytra usually completely deeply inserted under anterior pronotal margin. covering abdomen, striate or striate-punctate. Procoxae Mandibles moderately large. Antennae 11-segmented, transverse, globose, their cavities open or closed; with a large 3-segmented club. Pronotum strongly mesocoxae globose, broadly separate, open; metacoxae transverse and emarginate apically; posterior angles transverse, broadly separate. Legs with femora swollen. tightly join-ing angular humeri of elytra in lateral view, with lateral narrow portion of pronotum and elytra SYNOPSIS. In New Zealand, laemophloeids are represen- explanate. Scutellum Iarge and more or less triangular. ted by 3 adventive cosmopolitan species of Cryptolestes, 2 Elytra broadly rounded. Procoxae convex and trans- native species (1 undescribed) in Microbrontes, and a verse, their cavities open. Tarsi 5-5-5-segmented in third, apparently undescribed and tentatively assigned to both sexes, with the 4th segment reduced and the 3rd Notolaemus. Microbrontes lineatus is known from several lobed (Phalacrus), or 5-5-5 in females or 5-5-4 in males, localities in the North Island and from the Three Kings. with the 4th segment normally developed and the 3rd not Adults of Cryptolestes pusilloides have been collected in lobed (Cyclaxyra). Auckland, and adults of C. ferrugineus are known from several widespread localities. SYNOPSIS. There are 2 native phalacrids in New Zea- Most laemophloeids usually feed on moulds or on the land, with possibly a third undescribed, all in the endemic spores and stromata of ascomycete fungi (Lawrence & subfamily CYC A Y I AE and an adventive Australian Britton 1994), and live under bark of trees, but species of subspecies in A AC I AE known from Auckland and Cryptolestes are pests of stored products – see Booth et al. Rotorua (Kuschel 1990, Thompson & Marshall 1980). (1990), Bousquet (1990), and Harney (1993). Adults of Cyclaxyra impressa ranges from the Nelson area to the Microbrontes lineatus have been collected from ferns at Muttonbird (Titi) Islands off Stewart Island; C. politula is night, dead twigs of pohutukawa (Metrosideros excelsa), known from a few records on both main islands. and nests of Turdus sp. Cryptolestes pusilloides is known Adults of Phalacrus uniformis frigoricola have been from stored maize and wheat, and an adult specimen was found in rust fungus galls on an introduced wattle (Acacia collected in winter from a starling (Sturnus vulgaris) nest mearnsii). Adults and larvae of Cyclaxyra impressa occur in a bird box; C. ferrugineus has been found in stored in sooty mould growing on the honeydew of scale insects cereals, prunes, garlic cloves, and a drying kiln. on the trunks of mountain beech and red beech (Notho- fagus). Very little is known about C. politula, some RANGE. North ®, South ®, Stewart O records indicating the same habitat as for C. impressa, plus Kermadecs ®, Three Kings ®, Chathams O, Snares O moss on branches of a stunted Senecio sp. which also Aucklands O, Campbell O, Antipodes O, Bounties O frequently bears sooty mould.
-49.- 9- The biology of phalacrid beetles is reviewed by Steiner and pipit (Anthus novaeseelandiae) (Watt 1980b). Adults (1984). of Neocercus electus are known to occur in damp native forests in the northern South Island (Watt 1980b). RANGE. North ®, South ®, Stewart Kermadecs O, Three Kings O, Chathams O, Snares O RANGE. North ®, South ®, Stewart O Aucklands O, Campbell O, Antipodes O, Bounties O Kermadecs O, Three Kings ®, Chathams ®, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O EXAMPLE. CYCLAXYRINAE - Cyclaxyra impressa Broun, 1915 (Fig. 196). EXAMPLE. Zeonidicola chathamensis Watt, 1980 (Fig. 197). REMARKS. The systematic position of Cyclaxyra is controversial, and is under review by R.A. Crowson and REMARKS. It seems likely that cavognathids will be by S.A. Slipinski (Lawrence & Newton 1995). found in nests of other bird species. Suggested coloquial name: shining fungus beetles. Suggested colloquial name: birds' nest beetles. SELECTED REFERENCES. Crowson (1973), Pakaluk SELECTED REFERENCES. Pakaluk et al. (1994), et al. (1994), S n Gupta & Crowson (1969b), Watt Steiner (1984), Thompson & Marshall (1980). (1980b).
[49] Family CAVOGNATHIDAE [50] Family CRYPTOPHAGIDAE Fig. 197 [assisted by R. Leschen] Length 2.5-4.0 mm Tarsal formula 5-5-5 Fig. 198
Length 1-3.5 mm Tarsal formula 5-5-5 or (males DIAGNOSIS. Body narrowly elongate, broadly curved of Cryptophaginae) 5-5-4 laterally, somewhat depressed, moderately glossy; dorsal surface with decumbent pubescence; punctation DIAGNOSIS. Body oval to elongate oval, variously irregular; colour brown to reddish-brown, often with yellow-brown or red-brown to dark brown, sometimes paler appendages. Head narrower than pronotum; frons bicoloured, with appendages usually paler, often clothed sometimes with impressions; frontoclypeal suture ab- with silky hairs (decumbent and/or erect), rarely glob- sent; gular region with longitudinal grooves. Mandibles ose and appearing glabrous, convex to slightly flattened. each with 1 or 2 cavities opening outwards. Antennae Head narrower than pronotum, without a frontoclypeal 11-segmented, with enlarged scape and with a 3-seg- suture. Antennal insertions exposed, approximated or mented, elongate and loosely formed club. Pronotum well separated. Antennae 11-segmented, with a loose 3- simple, approximately square or rectangular in outline, segmented club (exception: 2-segmented in Picrotus). with margins broadly arched and angles rounded; lateral Eyes small. Pronotum subquadrate or rounded, with carinae indistinct; disc with a smooth, raised, median lateral carinae; lateral margins occasionally serrate impunctate area near base and with a shallow, punctate (introduced Cryptophagus); base often with a narrow, depression on either side. Elytra with anterior angles variously formed depression; prosternal process moderately (shoulders) slightly produced, and base concave; epi- broad, overlapping mesosternum; pronotal base usually pleura distinct in basal third; sutural flanges widened narrower than elytral base (usually as broad as in apically (Fig. 93), creating a small gap; striae absent. Mycetophagidae [59], or only slightly narrower). Elytra Procoxae slightly globose and transverse, their cavities subparallel or broadly arched laterally, rounded post- narrowly open behind. Legs stout. Protarsi and meso- eriorly; punctation confused, the punctures not in rows; tarsi with 3 basalmost segments expanded in males. sutural flanges widened apically, forming a smalI gap; SYNOPSIS. The cavognathids comprise 4 genera, all in epipleura incomplete, visible in basal third. Procoxae the Southern Hemisphere. Watt (1982a) listed 2 described globose, with cavities open behind. Abdomen with 5 native species of Zeonidicola and 1 of Neocercus for New ventrites, the basal one longest. Zealand. NZAC now has an additional 1 or perhaps 2 undescribed species of Zeonidicola. SYNOPSIS. Watt (1982a) reported 23 native and 4 ad- Adults and larvae of Zeonidicola dumbletoni have been ventive species of cryptophagid in New Zealand. There are found in nests of spotted shag (Stictocarbo punctatus). at present 18 native and 12 adventive species housed in Adults of Z. chathamensis have been extracted from nest NZAC, belonging to some 12 genera in 2 subfamilies, debris of sooty shearwater (Puffinus griseus), fairy prion CRYPTOPHAGINAE and ATOMARIINAE. (Pachyptila turtur), giant petrel (Macronectes giganteus), Cryptophagids in general feed on hyphae, conidia, and northern royal albatross (Diomedea epomophora sanfordi), spores of fungi, including moulds, in leaf litter, forest litter
-50- (e.g., beech forest), vegetable debris, rotten wood, bird notha) nd d t n t p n t t tr (irregularly punctate and insect nests, and under bark (Leschen 1996). Cosmo- in Loberonotha). r x l b th r v t p n politan species found in stored products feed only on r l d behind. r th th nt r d d; mould, their presence indicating poor storage conditions; nt nd 3 th t l b Bousquet (1990) reviewed stored products pests. Kuschel (1990) recorded 13 species from Auckland, collected from SYNOPSIS. Watt (l982a) recorded 8 native languriids heaps of garden prunings, lawn clippings, compost, hen- for New Zealand, 7 of them represented in NZAC. Our house straw, paddocks, leaf litter, Phellinus fungus, fresh- present estimate is 7 nominal species plus 1 undescribed; cut branches of various trees and shrubs, and foliage of these are placed in 5 genera representing 2 subfamilies — tree ferns; some were associated with sedges and galls. E OSCE I AE and C Y O I I AE Adults and larvae of Hapalips prolixus occur under RANGE. North ® South ® Stewart dead leaf bases of nikau palm (Rhopalostylis sapida) and Kermadecs O Three Kings O Chathams ®, Snares O in decayed tree-fern fronds. Adults of Loberus depressus Aucklands O Campbell O Antipodes O Bounties O have been collected by beating foliage of cabbage tree (Cordyline australis), pohutukawa (Metrosideros excelsa), EXAMPLE. C Y O AGI AE — Thortus ovalis Broun, and from tea tree blossom. Watt (1982b) recorded adults 1886 (Fig. 198). of L. nitens beaten from Coprosma repens, Carmichaelia sp., M l t r ίfl r and Pittosporum crassifolium. REMARKS. Cryptophagids are poorly known in the Adults of Loberonotha have been collected by beating Southern Hemisphere, and those placed in the Crypto- various vegetation, from litter, and from flowers of phaginae (Cryptosomatulini) are in need of revision Traversia. Adults of Cathartocryptus maculosus, our sole (Leschen & Löbl 1995). cryptophiline, have been found under bark of dead Suggested colloquial name: silken fungus beetles. Beilschmiedia tawa and on foliage of Coprosma robusta.
SELECTED REFERENCES. Leschen (1996), Pakaluk RANGE. North ®, South ®, Stewart ® et al. (1994). Kermadecs O Three Kings O Chathams O Snares O Aucklands O Campbell O Antipodes O Bounties O
[51] l A GU II AE EXAMPLE. E OSCE I AE — p l p prolixus (Sharp, [assisted by R. Leschen] 1876) (Fig. 199). Fig. 199 REMARKS. As at present defined, the family Languriidae Length 2-6 mm Tarsal formula 5-5-5 includes several genera formerly placed in Cryptophagidae This group is heterogeneous in appearance, and its classi- [50] and Erotylidae [52] (Lawrence & Newton 1995). fication is not clear. Some members are similar to Cryp- Suggested colloquial name: slender beetles. tophagidae [50], but are distinct in having the first ventrite less than twice as long as the second (cf. more than twice SELECTED REFERENCES. Pakaluk et al. (1994), S n as long in Cryptophagidae), posterior elytral sutural gap Gupta (1968), S n Gupta & Crowson (1969a, 1971). absent (cf. present), epipleura complete, hind wing with 4 anal veins (cf. usually 5) and a closed radial cell, usually a transverse groove on the anterior part of the gular region, [5 ] l E O Y I AE and 5-5-5-segmented tarsi in both sexes (Atomariinae of Fig. 200 Cryptophagidae have the same tarsal formula). For dis- Length 4-8 mm Tarsal formula 5-5-5 cussion see Sen Gupta (1968), Sen Gupta & Crowson (1969a, 1971), and Lawrence & Britton (1994). DIAGNOSIS. d l n t v l d r t l nv x DIAGNOSIS. d d r t l t tr n l l n t b t th l htl fl tt n d pr n t ll l br n rr l v l t bp r ll l b l ndr l t fl t- r th p r n n p p b n tr n l t n d pubescent or glabrous, moderately to strongly l with punctation forming striae on elytra. Color- glossy, pale brown to almost black, rarely with pale ation usually dark brown to nearly black with reddish- maculae on elytra. d d r t l l r th brown appendages, rarely yellowish brown. d d pl r l f t d Ant nn th d t n t l n t 3- n rt d nt th r x; fr nt l p l t r nd t n t r nt d l b r n t th l t r l r n pl b nt Ant nn 11- nt d th 3- nt d slightly crenulate in Hapalips and Loberonotha, ll l b E pr n nt and coarsely faceted. r n t d t n tl n rr r th n l tr l b ; base sometimes th h rp l t r l r n nd nt r rl pr d d with 2 small depressions. El tr ft n l n t nd fr nt n l ; base of pronotum and elytra slightly sinuate. ll th lI d f n d p pl r (absent in Lobero- El tr tr t th p pl r pl t Prosternal pro- -51- cess broad. r x l b n rt d nt rt l r SYNOPSIS. Hudson (1934) recorded 7 species of both- d pr n th r v t l d Legs robust. r riderid from New Zealand. Five are represented in NZAC, th ll nt x pt th l t l b d nd t belonging to the subfamilies A OMMA I AE with 1 adven- b n th r pl r l l n t pr nt on tive species, and O I E I AE with 4 native species. A basal ventrite, which is 1.5× as long as ventrite 2. review of the world Bothriderini is provided by Slipinski et al. (1989). SYNOPSIS. Watt (1982a) reported 9 native species of All our bothriderids are predacious. Adults of the ad- erotylid. Seven species are represented in NZAC, belong- ventive hypogeal species Anommatus duodecimstriatus ing to 2 genera of AC I AE are recorded from rotten wood buried in earth, under Both adults and larvae are in general mycetophagous, deeply buried stones (Sen Gupta & Crowson 1973), and feeding within fruiting bodies of mushrooms and bracket from deep pitfall traps sealed off at the top (Kuschel 1990). fungi. Kuschel (1990) reported two species of Crypto- Adults of Ascetoderes paynteri from the Chatham Islands dacne from Auckland, collected from moist decayed wood have been collected in bush at night, suggesting a noct- and from wood mould of Vitex. We have seen specimens urnal habit; others were taken from rotting Plagianthus from dead, standing Corynocarpus laevigatus, from under logs. Adults of A. obsoletus are occasionally found under various logs, in the dead bole of a bat roosting tree, from bark of dead tree trunks. Piptopterus portentosus on Nothofagus, from Ganoderma on a dead standing tree, and from Grifola colensoi. RANGE. North ®, South ®, Stewart O Kermadecs O Three Kings O Chathams ® Snares O RANGE. North ®, South ®, Stewart ® Aucklands O Campbell O Antipodes O Bounties O Kermadecs O Three Kings O Chathams ®, Snares O Aucklands O Campbell O Antipodes O Bounties O EXAMPLES. ANOMMATINAE - Anommatus duodecim- striatus (Müller, 1821) (Fig. 201); BOTHRIDERINAE — EXAMPLE. Dacninae — Thallis polita White, 1846 (Fig. Ascetoderes obsoletus (Broun, 1895) (Fig. 202). 200).
REMARKS. Suggested colloquial name: pleasing fungus REMARKS. Adult bothriderids superficially resemble beetles. some Colydiidae [65], from which they may be distin- guished by having exposed antennal insertions. SELECTED REFERENCES. Pakaluk et al. (1994). Suggested colloquial name: none.
SELECTED REFERENCES. S n Gupta & Crowson [53] l O I E I AE (1973), Slipinski et al. (1989). Fig. 201, 202
Length 2-7 mm Tarsal formula 4 4 4 (Ascetoderes), 3-3-3 (Anommatus) [5 ] l CE Y O I AE (= CE Y I AE DIAGNOSIS. d l n t - ll pt l b l ndr l t th pr n t l htl fl tt n d l br Fig. 203 l htl l dark brown to black. d n rr r Length 2-3 mm Tarsal formula 4 4-4 th n pr n t Ant nn 11- nt d th b l nt nl r d nd th - nt d l b; n- DIAGNOSIS. d n rr l l n t bp r ll l rt n xp d E lI nd pr j t n r r d d (Philothermus) r br dl v l (Hypodacnella), moder- (Anommatus). r n t tr p z d l th fr nt n l ately convex to flattened, l br t r t ll l htl pr d d coarsely punctate, and often d r t l t tr n l l often with a metallic shine with medial impression(s); base usually distinctly nar- (Hypodacnella), pale brown or dark brown to black, with rower than elytral base. El tr th r nd d h ld r fine or coarse punctures. d th t n p rt ll br dl r t l t r ll broadest in basal half, with n l d b pr n t ; frontoclypeal suture present or longitudinal striae and coarse punctation; alternate in- absent (Philothermus). Ant nn 1 - r 11- nt d tervals costate at least apically, even intervals usually th 1- r - nt d l b (superficially appearing 3- smooth; p pl r pr nt r x l b th r segmented in Hypodacnella). r n t tr n v r r v t p n (Anommatus) r l d (Ascetoderes); l htl l n t distinctly margined laterally, often metacoxae globose, moderately to widely separated. Legs coarsely punctate; base broadly arcuate or sinuate in l n t t t r l nt pl Abd n th Hypodacnella. El tr ll tr t -p n t t Hind b l v ntr t l n t t l t l n th f l- wings often with anal cell closed. r x lI nd l n v ntr t b n d l b l r xt rn ll th r v t p n r l d;
-5 - metacoxae widely separated. Abdomen with segments DIAGNOSIS. MYCETAEINAE (Mycetaea, Fig. 204). Body freely articulated; 1st ventrite markedly longer than small (length 1.8-2.0 mm), ovate and slightly tapering 2nd ventrite; femoral lines usually present on 1st posteriorly, moderately convex, yellow, with long and ventrite and/or metasternum. Last ventrite with post- sparse, decumbent and erect hairs. Head partly erior edge crenulate in Philothermus. concealed by pronotum; frontoclypeal suture distinct. Antennae 11-segmented, with an elongate 3-segmented SYNOPSIS. Watt (1982a) reported 5 native and 1 adven- club. Pronotum transverse, almost as broad as elytral tive species of cerylonid from New Zealand. We now base; front angles slightly produced anteriorly; lateral recognise 5+ native species in NZAC, belonging to 2 margins sharply carinate, with an additional lengthwise genera in 2 subfamilies, EUXESTINAE and CERYLONINAE. ridge on either side. Elytra with punctures forming Cerylonids occur in rotten wood, leaf litter, forest rows. Procoxae small, hemispherical, narrowly separ- debris, and fungus-infested bark (Slipinski 1988), where ated by prosternal process, their cavities open. behind. they probably feed on fungal hyphae and spores (Law- Abdomen with 1st sternite as long as the following 2 rence & Britton 1994). Kuschel (1990) reported Hypodac- combined, or somewhat longer. nella rubriceps (as Hypodacne) from Auckland, collected HOLOPARAMECINAE (Holoparamecus, Fig. 205). Body in rich, moist humus under logs, in hollow trees, and in minute (length 1.0-1.6 mm), narrowly oval, round pos- decayed wood. We have seen specimens collected from teriorly, moderately convex, reddish brown, yellowish, forest litter, rotten logs, and fungus on a Beilschmiedia or brown, appearing almost glabrous but with short, log, and reared from Helichrysum coralloides. Species of inconspicuous decumbent hairs, strongly glossy. Head Philothermus occur under bark of native trees, or in rotten somewhat concealed basally by pronotum; frontoclypeal logs and forest litter, especially with mould. suture present. Antennae 11-segmented, with a large, 2-segmented club. Pronotum restricted behind to a RANGE. North ®, South ®, Stewart ® distinct `waist,' with a series of laterally elongated Kermadecs O, Three Kings O, Chathams ®, Snares O foveae near posterior margin forming a basal impression; base markedly narrower than elytral base; additional long- Aucklands O, Campbell O, Antipodes O, Bounties O itudinal carinae absent. Elytra oval, with 2 impressed lines along suture; punctate striae absent. Procoxae EXAMPLE. EUXESTINAE — Hypodacnella rubriceps (Reit- small, hemispherical, narrowly separated by prosternal ter, 1880) (Fig. 203). process, their cavities appearing narrowly closed be- hind. Abdomen with 1st ventrite elongate, as long as the REMARKS. Some cerylonids may be confused with following 3 combined. Colydiidae [65], from which they differ, besides other characters, in having the upper body surface glabrous or SYNOPSIS. Endomychids are represented in New Zea- microsetose and glossy (cf. either dull or roughly sculp- land by an adventive European species of MYCETAEINAE tured), the maxillary and labial palps geniculate (cf. never and at least 6 native species of HOLOPARAMECINAE, 3 of so), the 1st ventrite markedly longer than the 2nd, and all them in Holoparamecus and the remainder in an undet- 5 ventrites movably articulated (cf. 1st ventrite not mark- ermined genus. Watt (1969) classified Holoparamecus in edly longer, and 3 or 4 basalmost ventrites connate in subfamily Merophysiinae of the Corticariidae [58], and Colydiidae). Slipinski (1988) provided characteristics of considered Mycetaea lucidus to be a synonym of Holo- the genera of Cerylonidae. This group is in need of re- paramecus tenuis. A new classification is proposed by vision. Pakaluk et al. (1994) and Lawrence & Newton (1995). Suggested colloquial name: waxen beetles. Our holoparamecines occur in forest litter, organic litter probably associated with nest debris, wood mould, and SELECTED REFERENCES. Pakaluk et al. (1994), S n bird nest litter; some adults have been reared from bracket Gupta & Crowson (1973), Slipinski (1990). fungus growing on dead roots, and others from dead Coprosma robusta. Species of this group most likely feed mainly on moulds associated with decomposing organic [55] Family ENDOMYCHIDAE material. Kuschel (1990) reported 2 species of Holo- (incl. former MEROPHYSIIDAE) paramecus from Auckland, collected from logs, stumps Fig. 204, 205 covered with fruiting bodies of larger fungi, rotted wood, moulds inside hollow trees, humus under logs and at the Length 1-2.0 mm Tarsal formula 4 4 4 (Mycetaea), base of herbaceous plants, and stream-bed litter. 3-3-3 (Holoparamecus) New Zealand species of this family belong to two genera in RANGE. North ®, South ®, Stewart O separate subfamilies, which are more readily diagnosed Kermadecs O, Three Kings ®, Chathams O, Snares O separately, as follows. Aucklands O, Campbell O, Antipodes O, Bounties O
-53- EXAMPLE. MYCETAEINAE - M t subterranea (Fab- ture, and 6 are known to be well established. In general, ricius, 1 1 (Fig. ; HOLOPARAMECINAE - l p r - members of the subfamily Coccinellinae usually feed on t n Reitter, 1 79 (Fig. 205). aphids and homopterans, and occasionally on larvae of Chrysomelidae and other Coccinellidae, whereas the REMARKS. The higher classification of Endomychidae Scymninae feed on aphids, scale insects, mealybugs, was discussed by Lawrence (1982), Slipinski (1990), and whiteflies, and plant mites (Booth et al. 1990). Kuschel Slipinski & Pakaluk (1992). (1990) listed 26 coccinellid species from Auckland, most Suggested colloquial name: waisted mildew beetles. of them in genus Rhyzobius, and the majority collected from garden plants and foliage of shrubs and trees. Very common in Auckland is the introduced Australian species SELECTED REFERENCES. Pakaluk et al. (1994), Watt Halmus chalybeus, steel-blue in colour, and occurring in (1969). gardens and native bush. Collecting: sweeping and beating vegetation.
[56] Family COCCINELLIDAE RANGE. North ®, South ®, Stewart Fig. 206 Kermadecs ®, Three Kings ®, Chathams ®, Snares O Length 1-7 mm Tarsal formula 4-4-4 or 3-3-3 Aucklands ®, Campbell ®, Antipodes ®, Bounties O DIAGNOSIS. Body narrowly to broadly ovate or EXAMPLE. COCCI E I AE - C n ll l n n br - hemispherical, moderately to strongly convex, with cius, 179 (Fig. 206). ventral surface usually flat, glabrous or pubescent, slightly to strongly glossy or matt. Setation short and REMARKS. Species introduced for biological contol of decumbent or long and protruding, often yellowish or hemipteran pests are in Coccinella, Cryptolaemus, Exo- greyish. Coloration usually uniformly brown in smalI chomus, Halmus, Hippodamia, Illeis, Rhyzobius, Rodalia, species and bicoloured in larger ones, variously pale and Scymnus. The family is in need of taxonomic revision. brown, brown, yellowish, reddish brown, black, or met- Suggested colloquial name: ladybirds. allic blue, often with spots on pronotum and elytra or with a paler margin on pronotum and spots on elytra, the spots SELECTED REFERENCES. Bielawski (1976), Klaus- yellow, reddish yellow, reddish brown, or black. Head nitzer (1970), LeSage (1991 c), Pakaluk eta!. (1994), Pope partially or (rarely) almost entirely inserted into (1989), Read (1965), Sasaji (1968). prothorax; frontoclypeal suture absent. Maxillary palps with apical segment usually enlarged, securiform or oblong, and apex obliquely truncated. Antennae [57] Family CORYLOPHIDAE usu-ally 10- or 11-segmented, short, with a weak and Fig. 207 usually 3-segmented club. Pronotum transverse, usually Length 0.8-2 mm Tarsal formula 4-4-4 or 3-3-3 strongly convex; lateral margins entire; anterior margin often strongly emarginate; scutellum small to large. DIAGNOSIS. Body minute, oblong to broadly ovate or Elytra usually strongly rounded, with confused punc- nearly hemispherical, moderately to strongly convex, tation, never striate; lateral margins entire; epipleura sparsely pubescent or glabrous, often glossy. Coloration sometimes foveate to receive femora. Procoxae trans- pale brown, brown, yellowish with brown markings, red- verse, their cavities open. Tarsi pseudotrimerous or dish brown, or reddish yellow. Head small, completely trimerous, with segments 1 and 2 strongly produced concealed by expanded pronotum forming a hood beneath. Abdomen with 5 or 6 ventrites, the basalmost anteriorly (partially concealed in Orthoperus); fronto- 2 often connate; basal ventrite with impressed femoral clypeal suture absent. Antennae 9-11-segmented, with lines (Fig. 92). a 3-segmented club. Pronotum expanded anteriorly into a sharp margin over head, strongly transverse, SYNOPSIS. Watt (1982a) reported 27 described native broadly rounded; posterior angles sometimes strongly species of coccinellid and 14 adventive or introduced. produced posteriorly (Anisomeristes) or slightly conically There are at present 22 native species and about 18 elongate (Arthrolips). Elytra subparallel or narrowed representing introductions in NZAC, classified into some posteriorly and usually truncate, exposing pygidium. Pro- 19 genera in 4 subfamilies — COCCIDULINAE, SCYMNINAE, coxae transverse or globose, their cavities closed. Tarsi CHILOCORINAE, and COCCINELLINAE. with 2nd segment bilobed, 3rd segment vestigial and Most known adults and larvae are predators, feeding inconspicuous. Abdomen with 6 ventrites, the basal- generally on soft-bodied insects and mites. A considerable most Yery long and bearing subparallel femoral lines. number of exotic species have been introduced into New Zealand for biological control of insect pests in agricul- SYNOPSIS. Watt (1982a) recorded 5 native and 1 adven-
-54- tive species of corylophid in New Zealand, but the actual (exception: M l n phth l often with ridges and/or number is much higher, with many species undescribed. depressions, or evenly convex and punctate, sometimes They belong to 8 genera in 3 subfamilies — CORYLOPHINAE, with a single basal depression; lateral margins smooth or SERICODERINAE, and PARAMULINAE. Kuschel (1990) repor- with fine dentition (e.g., C rt r Elytra elongate ted 20 species in 8 genera from Lynfield, Auckland alone. oval, broadest at middle, usually much broader than Corylophids are known to feed on fungi, and may be pronotum, or oval, sometimes swollen (M l n phth l found in association with decaying plant material such as with more or less distinct punctate striae. Procoxae pro- forest litter, organic litter generally, compost heaps, birds' jecting, their cavities closed behind. Tarsal segments nests, grass cuttings, mouldy hay, rotten wood, dead simple. Abdomen with 5 or 6 ventrites, the basal one branches, litter in coastal cliff vegetation, and leaf litter at markedly elongate (much longer than the 2 following the base of sedges and flax. Some New Zealand species ventrites combined). have been collected from under bark, on mossy vegetation at night, on logs overgrown with moss, on moss and liver- SYNOPSIS. Hudson (1934) recorded 57 nominal species worts on damp slopes near streams, on foliage of native of corticariid for New Zealand. Watt (1982a) recorded 40 shrubs, occasionally on flowers, and on bracket fungi. native and 11 adventive species, and some 53 native Sifting organic material and processing it through species and 11 adventive are now represented in NZAC. Berlese funnels is the most efficient method of collecting; Our corticariid fauna is rich, with many undescribed for details see Kuschel (1990). species, belonging to some 12 genera in 2 subfamilies, LATHRIDIINAE and CO ICA ΙI AE K h l (199 l t d 3 RANGE. North ®, South ®, Stewart ® species in 9 genera from Auckland. Kermadecs ®, Three Kings ® Chathams ®, Snares O Lathridiines are subglabrous, with separated procoxae, Aucklands O, Campbell ®, Antipodes ®, Bounties O often with carinae on the pronotum and elytra, and waxy exudate on various parts of the body; corticariines are EXAMPLE. S r d r n — Anisomeristes sharpi Mat- usually setose, with contiguous procoxae, lacking carinae thews, 1 (Fig. 7 on the pronotum and elytra, and with no waxy exudate. Adults and larvae feed on fungal spores. They occur in REMARKS. Broun (e.g., 1886, 1893a) habitually treated leaf litter, decaying trunks and stumps, wood mould, cut branches, foliage of various plants, bracket fungi, moss corylophids as Coccinellidae [56]. Corylophids are poorly and liverworts on tree trunks, hollows of old trees, bush known, and the subfamily classification is not settled, floor litter, epiphytes, lawn clippings, and garden com- although Lawrence & Newton (1995) have listed 4 sub- post. Some cosmopolitan species are associated with families in the world fauna. The group is in need of taxo- recorded from a wide nomic revision. stored products, and Ar d n d f r variety of mouldy vegetables, is now established in New Suggested colloquial name: hooded beetles. Zealand. For other cosmopolitan species of economic importance see Booth t l (1990). For details on the SELECTED REFERENCES. Endrödy-Younga (1964), bionomics of species occurring in Auckland, see Kuschel Gressitt & Samuelson (1964b), Pakaluk t l (1994). (1990).
RANGE. North ®, South ®, Stewart ® [58] Family CORTICARIIDAE Kermadecs ®, Three Kings ®, Chathams ®, Snares O (= LATHRIDIIDAE = LATRIDIIDAE) Aucklands ®, Campbell ®, Antipodes O, Bounties O Fig. 208 LATHRIDIINAE - Broun, Length 0.8-2.2 mm Tarsal formula 3-3-3 EXAMPLE. En v p (1893) (Fig. 208). DIAGNOSIS. Minute oblong beetles, with pronotum usually markedly narrower than elytra (as broad as REMARKS. The name Corticariidae has priority over the elytral base, or nearly so in M t phth l (= th t - universally used Lathridiidae for this family (Pakaluk t n Body surface glabrous or finely tomentose, often l 1994). First revisions of New Zealand corticariid spec- covered by a waxy exudate, moderately glossy to matt, ies were published by Belon (1884, 1897). Watt (1969) smooth and punctate or with ridges, tubercles, and/or provided a generic key to New Zealand Corticariidae, depressions. Head with clypeus laterally expanded including l p r now referred to Endomychidae in front of antennal insertions; frontoclypeal suture pres- [55]. ent. Antennae 11-segmented, with a 2- or 3-segmented Suggested colloquial name: mildew beetles. club; scape usually enlarged, often globular. Eyes small but protuberant, sometimes reduced, coarsely faceted. SELECTED REFERENCES. Belon (1884, 1897), Gressitt Pronotum usually distinctly narrower than elytra at base & Samuelson (1964a), Pakaluk t l ( 4 Watt (1969).
-55- S p rf l E E IO OI EA (HETEROMERA) [ ] l A C EOC Y ICI AE Members of this group are distinct in having tarsi 5-5-4- Fig. 210 segmented in both sexes, rarely 4-4-4 in a few lineages, Length 2.7-3.5 mm Tarsal formula 5-5-4 trochanterofemoral attachment strongly oblique (Fig. 59), procoxae tending to be conical and projecting, hind wings DIAGNOSIS. d n f r l v l l htl fl tt n d with a reduced number of veins (4 or fewer) behind the dark brown to almost black, with short pubescence. d media posterior (MP), aedeagus heteromeroid with the p rt ll r tr t d nt pr th r x br dl r t tegmen incomplete ventrally, and in certain characteristics nt r rl ; fr nt l p l t r d t n t M x ll r of the larvae (Lawrence & Britton 1991, 1994). p lp th p l nt nl r d nd tr n t Ant nn 11- nt d th 3- nt d l b E r l f t d r n t tr n v r th h rp [59] l MYCE O AGI AE l t r l r n slightly emarginate anteriorly and in- Fig. 209 significantly sinuate basally. Elytra arcuate laterally, with
Length 1.5-3 mm Tarsal formula 4-4-4 (females), confused punctation. r x l b p r t d b 3-4-4 (males) pr t rn l pr h h r d ll xp nd d p - ll r l nt pl Abd n th 5 v nt- IAG OSIS d br dl v l h t fl tt n d r t th f r t nn t (3 nn t n n br n d v nl p b nt d r t l l p n t t yellowish [ ] ; b l v ntr t l htl l n r th n nd v ntr t to dark brown to black, sometimes with pale or dark markings on elytra and/or pronotum. d n rr r SYNOPSIS. There is a single adventive species of arch- th n pr n t ; nt nn l n rt n xp d M x ll r eocrypticid, Archeocrypticus topali, in New Zealand. p lp th l t nt nl r d Ant nn 11- Adults and larvae are reported from leaf litter and some nt d ll th 3- nt d l b E polypore fungi with softer fruiting bodies (Lawrence & r l f t d r n t tr n v r ; d r t Britton 1994). Our species has been recorded from gar- t th n t rr t n thl nt n dens, parks, litter in beech forest, under stones (rarely), th d f l tr ; base as wide as elytra. Scutellum under seaweed on beaches, moss on banks, nests of gulls transverse. El tr br dl v l punctate. r x (Larus sp.), on mummified apples, and under mature tr n v r th r v t p n Legs well developed. cabbages. In all these habitats it is probably associated with moulds or other fungi. Many adults and larvae have SYNOPSIS. Watt (1982a) reported 12 native and 1 ad- been collected in pitfall traps in pastures around Nelson. ventive species of mycetophagid for New Zealand. These are in 3 genera of subfamily MYCETOPHAGINAE. Kuschel RANGE. North ®, South ®, Stewart O (1990) recorded 5 native species and 3 adventive from Kermadecs O Three Kings O Chathams O Snares O Lynfield, Auckland. Aucklands O Campbell O Antipodes O Bounties O Adults and larvae may be found on or in fungi and organic material infested with fungi. Some species have EXAMPLE. Archeocrypticus topali Kaszab, 1964 (Fig. been collected by beating or sweeping foliage of native 210). shrubs. The adventive species have been found in a sheep carcass, coarse prunings, a starling nest, bumble bee nests, REMARKS. Suggested colloquial name: ancient fungus lawn clippings, compost, trees, shrubs, and a rotting seed beetles. head of Cynara scolymus (Kuschel 1990). SELECTED REFERENCES. Kaszab (1981, 1984), Law- RANGE. North ® South ®, Stewart O rence (1994), Watt (1974a). Kermadecs O Three Kings ®, Chathams ®, Snares O Aucklands O Campbell O Antipodes O Bounties O [ 1] l CII AE
EXAMPLE. MYCETOPHAGINAE — Triphyllus hispidellus Fig. 211
(Broun, 1880) (Fig. 209). Length 1-3 mm Tarsal formula 4 4 4 REMARKS. Mycetophagids are poorly known in New DIAGNOSIS. d l n t b l ndr l r n rr - Zealand. The European concept of Triphyllus applied to l v t nd d r t l nv x yellowish, brownish, our native species needs clarification. reddish brown, dark brown, or greenish, uniform in colour Suggested colloquial name: hairy fungus beetles. or with yellowish or brownish elytral spots sometimes of irregular shape, l br nd tr n l l r d- SELECTED REFERENCES. Lawrence (1987), Leschen r t l l nd th h rt d b nt h r nd/ r & Lawrence (1991), Parsons (1975). br tl d d fl x d th d t n t fr nt l p l
—5 — r d th n l t f r n pl t r l n t t l t n t r nd t t r nd t b r l r h rn Maxillae with reduced lobes and p n lt t nt l b d Abdominal ventrites mov- fusiform palps. Ant nn -1 - nt d th ll able (not fused). 3- nt d l l b (Fig. 9) (cf. a similar 3-seg- mented compact club in Curculionidae: Scolytinae [82]). SYNOPSIS. Watt (1982a) reported 38 native species of r n t l r p rt ll r nt r l v r n h d melandryid for New Zealand. The species are assigned to (Fig. 9); base often approximately straight; l t r l d 9 genera, 5 in subfamily ME A YI AE and the remainder n pl t El tr l n t ll bp r ll l th of uncertain affiliation. Kuschel (1990) recorded 8 species p n t r nf d r n r r x tr n v r in 7 genera from Auckland. pr j t n th r v t p n b ft n th p n Adult melandryids occur in rotten wood, logs, organic n t r d Abdomen with 5 ventrites; basal ventrite litter including leaf litter, cut branches, flowers, at the usually with a setose fovea in male. base of bush-floor sedges, and on forest foliage. Adults are very agile, and are capable of jumping; some have been SYNOPSIS. Watt (1982a) recorded 20 native species of collected using interception traps. Larvae inhabit hard ciid for New Zealand. These belong to 5 genera of substrates, such as bracket fungi or wood (Lawrence & subfamily CII AE Kuschel (1990) recorded 17 species Newton 1995). from Lynfield, Auckland, in 5 genera. Adults and larvae are typically mycetophagous in fruit- RANGE. North ®, South ®, Stewart ing bodies of shelf fungi (Polyporaceae). Some occasion- Kermadecs O Three Kings ®, Chathams ®, Snares ally occur in forest litter, dead wood, or cut branches. Aucklands ®, Campbell ®, Antipodes ®, Bounties O
RANGE. North ®, South ®, Stewart ® EXAMPLE. ME A YI AE- Hylobia nubeculosa Broun, Kermadecs O Three Kings ®, Chathams ®, Snares O 1886 (Fig. 212). Aucklands O Campbell O Antipodes O Bounties O REMARKS. Suggested colloquial name: leaping beetles. EXAMPLE. CII AE - Cis zeelandicus Reitter, 1880 (Fig. 211). SELECTED REFERENCES. Crowson (1966b), Gressitt & Samuelson (1964a), Viedma (1966). REMARKS. Suggested colloquial name: shelf fungus beetles. [ 3] l MO E I AE SE EC E REFERENCES. Abdullah (1973), Lawrence Fig. 213 (1971), Zimmerman (1938). Length 4-17 mm Tarsal formula 5-5-4 IAG OSIS d d d- h p d t p r d p t r - [ ] l ME A YI AE rl l t r ll pr d h r t r t ll h p d Fig. 212 n l t r l v th th l t bd n l nt Length 1.5-12 mm Tarsal formula 5-5-4 pr d d nd f r n n t pr xt nd n b nd l tr ( ft n xtr l l n t ; rf th IAG OSIS d l n t ppr x t l d - f n d b nt p b n ; coloration brown with h p d t p r n p t r rl nv x d r ll nd darker markings on pronotum, uniformly black, or black v ntr ll th f n nd d b nt p b n finely with small white spots on elytra and often on lateral parts or coarsely punctate; coloration uniformly dark brown to of sternites. d d fl x d fl t v ntr ll nd n l n black, pale brown, or yellowish orange, sometimes with pr t rn ( n r t n p t n br ptl n tr t d yellow, orange, or brown markings on elytra and pro- b h nd nd f r n n rr n M x ll r notum. d d fl x d d pl n rt d nt pr th r x p lp th l t nt xp nd d Ant nn 11- n t n tr t d b h nd M x ll r p lp th l t - nt d f l f r r l htl rr t Eyes entire. nt nl r d ft n r f r Ant nn ll r n t br d l tr r l htl br d r f l f r r n r t b t th t d t n t l b Eyes transverse, sinuate basally and th pl t l t r l entire, not pubescent. r n t th l t r l r n r n El tr t p r n p t r rl r x pr j - n pl t nt r rl sometimes with small longitudinal t n th r v t p n; t x nl r d pl t - depressions. El tr t p r n p t r rl sometimes l b ft n th b- h p d p n with impressed longitudinal lines subparallel to suture. r x pr j t n th r v t p n behind. Legs SYNOPSIS. Watt (1982a) recorded 5 native species of l n b l p r ft n b- h p d xtr l l n- mordellid for New Zealand. Kuschel (1990) listed 6 t n t t b (Fig. 84). r th b l nt species, including 1 adventive, in 4 genera from Lynfield,
—57— A l nd n r r n bf l MO E I AE nd n f r b d b tl K h l (199 r p rt d th r nd r r n n d All n p br t b pr d t r n l rv f O n Ad lt h v ft n b n f nd n fl r f n h rt (C r b d h l rv r n n t b ( pt p r p r r rt (S n j b nt ph p r t d f t r n pt r b ll f r fl r f l f n t v hr b nd tr C l pt r (C r b d nd l tt d nd th r f r t l tt r nd r rv r n r tt n d r d v l p nt nv lv b th xt rn l nd nt rn l f d n t f v r p t r l pl nt ( n t (S l nd r 1991 Unl th r rh p ph r d th n l rv f h p t n n r f r pp r ntl fr -l v n C ll t n p n r b t n f l f tr hr b pr d t r f l rv f r n pl (C r b d ( d n nd l r v t t n r fr nt r pt n tr p 193
A GE rth ® S th ® St rt A GE rth ® S th ® St rt O K r d O hr K n O Ch th ® Sn r O K r d O hr K n O Ch th O Sn r O A l nd O C pb ll O Ant p d O nt O A l nd O C pb ll O Ant p d O nt O
E AM E MO E I AE — M rd ll nt r t Wh t E AM E E ECA OMI AE— h p t n l br Sh rp 1 ( 13 1 7 ( 1
EMA KS l nd rd ll d r rr ntl EMA KS h r l t n h p f r rh p ph r d n r b n t d d b r M E r n l f G n It l n th rld nt xt r r nv t t n S t d ll l n p nt l b tl S t d ll l n ntl r d b tl
SE EC E E E E CES tt n (199 Er h SE EC E E E E CES S l nd r (1991 (195 r n l (19 Odn (1991 [ 5] l CO Y II AE [ ] l I I O I AE 15 1 1 n th 1-19 r l f r l - - n th - r l f r l 5-5- IAG OSIS d br dl v l r v t t n rr l IAG OSIS d l n t r r l l t r ll bp r ll l t xtr l l n t ( pr d t p r n p t r rl b t l n n nv x t n t t fl tt n d ft n th r d pr t b r- bd n l p n f n l p b nt th p b n d r n r rn nt t n d r n l r ll br n r ll h nd d b nt; l r t n p l br n t t bl t v r t d th p l r r d r r r - d r br n th h d t bl nd l tr ll n ll tt th v r p b n d b nt p l r d d fl x d ( n n t r l p t n br ptl r r t r th br tl r l r r l l nd n tr t d b h nd t f r n Ant nn th b l br d ll p rt ll r tr t d nt th - l t 7 nt fl b ll t n l ( 33 E r x ft n th ll d v l p d fr nt l p l r d ll r n t r r l dn - h p d nd h h b xp nd d f r rd ; nt nn l n rt n p rtl rr nd n b f nt nn r n t tr p z- ll n l d M x ll r p lp th l t nt d l n t b ll El tr v r n bd n nt r l ll Ant nn 1 - r 11- nt d n r t th r h rt n d nd xp n t l t bd n l p x th - r 3- nt d l b r n t ll tr n v r l n t d n l r d b t n t tr t r x pr j t n n ll dr t r l n t ll xpl n t th r v t p n b h nd l nd r b l p r l t r ll th l t r l r n pl t nd ft n n n- rr t r l nt pl ; t r l l r n l t r d nt t t xp nd d l t r ll t th d ( 7 nd th p ll pr d d fr nt n l ; d ll r v d r n t nd/ r t b r l t El tr th r d SY O SIS d n (193 l t d 5 n t v p f nd/ r t b r l nd r f p n t t n r x rh p ph r d f r l nd b t W tt (19 r rd d l b l r th r v t p n r l d r l nt nl h p r pl d n 3 n r f bf l pl Abd n th 5 v ntr t th b l 3 fr r E ECA OMI AE b t th t t f 1 n n rt n nd nn t t h v t b n n d K h l (199 r rd d 1 p fr nf ld A l nd SY O SIS W tt (19 r p rt d 19 n t v p f Ad lt r f nd n r bb v t t n n d d d l d d f r l nd h r r 133 p r pr - nd n f l f n t v hr b th n r rd fr th nt d n AC b l n n t n r n b-
— 5 — families, PYCNOMERINAE and COLYDIINAE. The status of tuberculate, with complete epipleura. Procoxal cavities some genera needs revision. Kuschel (1990) recorded 24 closed behind by lateral extensions of prosternal species in 14 genera from Lynfield, Auckland. intercoxal process (Fig. 39) (in Tenebrionidae [68] Pycnomerines are distinct by having partly closed closed by mesal extensions of pronotal hypomera — Fig. procoxal cavities, a laterally expanded prosternal process, 38). Tarsal segments simple. Abdomen with 3 basal and widely separated metacoxae, and by features of their ventrites connate. larvae; colydiines are distinct by having procoxal cavities either open or closed by projections of the notum, SYNOPSIS. Watt (1982a) reported 20 native species of moderately widely separated metacoxae, and by larval ulodid (as Zopheridae) for New Zealand. Fourteen species characteristics (Lawrence & Britton 1991). are represented in NZAC, belonging to 4 genera, of which Adults may be found under bark, on standing dead tree one is confined to New Zealand. trunks, in rotten wood, cut branches, leaf litter, forest litter, Adults and larvae of Brouniphylax squamiger occur in and on various trees and shrubs. Most species are believed dead bracket fungi (Fomes), and adults alone are some- to be mycetophagous, but Lawrence & Britton (1994) times found under logs. Both stages of Syrphetodes are reported certain New Guinean species as being predators found under loose bark of standing dead trees or logs and of platypodine weevils. Larvae are generally taken from in rotten branches; adults also are found on the underside rotten wood. of logs. Kuschel (1990) reported B. squamiger in dead Polyporaceae on decaying logs from Lynfield, Auckland. RANGE. North ® South ® Stewart Kermadecs ®, Three Kings ®, Chathams ®, Snares O RANGE. North ®, South ®, Stewart O Aucklands ® Campbell ® Antipodes O, Bounties O Kermadecs O, Three Kings ®, Chathams O, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O EXAMPLES. COLYDIINAE — Pristoderus antarcticus (White, 1846) (Fig. 215), Rhizonium antiquum Sharp, EXAMPLE. Syrphetodes marginatus Pascoe, 1875 (Fig. 1876 (Fig. 216). 217).
REMARKS. The placement of Rhizonium antiquum in REMARKS. Ulodids are superficially similar to tenebri- Colydiidae is uncertain, and should be treated as a tem- onids [68], from which they can be readily separated by porary arrangement. Lawrence (1994) has shown that having exposed antennal insertions, an apically expanded Pycnomerinae are more closely related to Monommatidae prosternal process, and in larval features (clypeus fused to and Zopheridae (sensu Lawrence & Newton 1995) than frons, and frontoclypeal suture obliterated). In tenebrionid they are to Colydiinae. larvae the clypeus is always distinct from the frons. For New Zealand's Colydiidae are under investigation by comparisons of Ulodidae, Chalcodryidae [67], and Zoph- Dr M.A. Ivie (Montana State University, Bozeman, eridae see Watt (1974a), Doyen & Lawrence (1979), U.S.A.) and by Dr S.A. Slipinski (Polish Academy of Lawrence (1994), and Lawrence & Newton (1995). For a Science, Warsaw, Poland). key to all genera of Ulodidae see Lawrence (1994). Suggested colloquial name: rough mould beetles. Suggested colloquial name: false darkling beetles.
SELECTED REFERENCES. Sharp (1876c), Slipinski & SELECTED REFERENCES. Doyen & Lawrence (1979), Burakowski (1988), Wollaston (1873). Lawrence (1994), Watt (1974a).
[66] Family ULODIDAE [67] Family CHALCODRYIDAE (formerly as ZOPHERIDAE) Fig. 218 Fig. 217 Length 5-22 mm Tarsal formula 5-5-4 Length 4-20 mm Tarsal formula 5-5-4 DIAGNOSIS. Body fairly soft, loosely articulated, nar- DIAGNOSIS. Body broadly oval, slightly depressed, rowly elongate, subparallel, moderately convex, glossy, brown, matt (exception: Arthopus, glabrous and strongly pale to dark green or brownish to dark brown, often glossy); dorsal surface with decumbent and/or erect with metallic reflections, and sometimes with patches of pubescence, bristles, or scales usually forming a pat- yellowish pubescence; surface punctate. Head prognath- tern. Head partially retracted into prothorax; antennal ous, slightly narrowed behind eyes, with a broad neck; insertions exposed. Antennae 11-segmented, filiform, antennal insertions exposed. Maxillary palp with last with at most a weak club. Eyes almost circular. Pro- segment enlarged, securiform. Antennae 11-segmented, notum narrower at base than elytra, often with front filiform, sometimes with last 3 segments slightly larger angles produced anteriorly. Elytra usually broad, often but not forming a distinct club. Pronotum transverse,
—59— r t n l r th d xpl n t El tr bp r ll l [ ] l E E IO I AE without clearly defined striae but with irregular rows of Fig. 219-221 punctures; p pl r th r n xt nd n l t t p x r x pr j t n th r v t l r tr n - Length 2.9-18 mm Tarsal formula 5-5-4 or v r l d behind partly by inward extensions of (Archaeoglenes) 4-4-4 propleura and partly by lateral extensions of narrow Adult tenebrionids are highly variable in form, and some intercoxal process; mesocoxal cavities narrowly separated, are superficially similar to members of other families, open laterally. Legs long and slender. Tarsal segments e.g., Carabidae [2], Scarabaeidae [15], Ulodidae [66], and elongate, not lobed. Abd n th 3 r b l v ntr t Chrysomelidae [77]. However, they are distinct in the l nn t following combination of characters.
SYNOPSIS. Chalcodryids are the only endemic family in DIAGNOSIS. d ll bl r br n rarely pale, New Zealand, with 5 native species in 3 genera. A key to moderately glossy (rarely glossy metallic) to matt, ft n genera and species is provided by Watt (1974b), together b l br d n t h rpl n tr t d b h nd with a detailed description of this family. Distribution ; nt nn l n rt n nd r nth (Fig. 11), records suggest a northern limit to the south of Auckland. h h n l t l t b f p fr b v nd Watt (1974b) collected adults by beating branches ll xt nd b t nt r r r n f on covered with moss and/or lichen in cool, wet forests, which it usually encroaches. M x ll r p lp th l t especially of Nothofagus. Adults of Chalcodrya are en- nt ft n r f r Ant nn ll f l f r countered most frequently, and their larvae have been n r t d t n tl l bb d r bl ntl rr t E found in short refuge galleries in dead suppressed twigs or ll r n t r n t ll r n t r small branches on various trees; they can be collected at xpl n t l t r ll El tr ll n rr l v l r night from moss-covered branches (Watt 1974b). Larval bp r ll l th d t n t p pl r if striate then gut contents predominantly consist of lichen and moss, usually with 9 or fewer striae (exception: 10 striae in some with some fragments of mites and spiders (Watt 1974b). Lagriinae), often with a scutellary striole or with sutural striae diverging basally. nd n t r d d RANGE. North ®, South ®, Stewart r x v r bl n h p ft n pr j t n th r Kermadecs O Three Kings O Chathams O Snares O v t l t l l d b h nd b xt n n f Aucklands O Campbell O Antipodes O Bounties O pr n t l h p r but not by lateral extensions of prosternal intercoxal process as in Ulodidae [66] and EXAMPLE. Chalcodrya variegata Redtenbacher, 1868 Chalcodryidae [67], without exposed trochantins. r h- (Fig. 218). nt r h t r r d (e.g., Fig. 59). Legs variable in form and function, from stout and specialised for digging to REMARKS. Chalcodryids in many features are similar to slender for running. r l nt ll n t l b d Periomylopidae, Ulodidae, and Tenebrionidae. They may b l th l pl r p t n t (Alleculinae) but be distinguished from Tenebrionidae [68] in having the never appendiculate. Abd n th 5 v ntr t th f r t procoxal cavities partly closed by lateral extensions of the 3 nn t and often with intersegmental membrane prosternal intercoxal process, 4 connate abdominal stern- clearly visible between ventrites 3/4 and 4/5. ites (3 in Tenebrionidae), and exposed antennal insertions. Chalcodryids differ from Ulodidae [66] in their fairly soft SYNOPSIS. Tenebrionids are a major group, comprising and loosely articulated body (cf. compact and strongly approximately 15 000 described species worldwide. There sclerotised), narrow prosternal intercoxal process, and are 149 valid species in New Zealand, 10 of them adven- lack of concealed lateral procoxal extensions. According tive, in 36 genera, 16 tribes, and 8 subfamilies (Watt 1992) to Watt (1974b), chalcodryids differ from Periomylopidae - AG II AE O O I I AE IME II AE E E IO I AE in having concealed trochantins on the procoxae and A ECU I AE IA E I AE and COE OME O I AE Tenebri- having usually 4 connate abdominal sternites. onids constitute 3.3% of the total known species of Lawrence (1994) transferred the Tasmanian genus Coleoptera in New Zealand (Watt 1992). Kuschel (1990) Sirrhas—placed in Chalcodryidae by Watt (1974b)—to recorded 21 species in 15 genera from Lynfield, Auckland. the Periomylopidae, leaving Chalcodryidae restricted in The biology of New Zealand tenebrionids is recorded distribution to New Zealand. Lawrence (1994) discussed by Watt (1992). Many of our native species live in rotten the new concept of the family. wood (Aphtora, Menimus, Uloma, Ulomotypus, Zolodi- Suggested colloquial name: southern beech beetles ( nus), or in powdery, dry rotten wood (Mimopeus opac- Palma, in litt.) ulus). Archeoglenes occurs in leaf litter and under logs. Menimus inhabits dead fruiting bodies of large woody SELECTED REFERENCES. Lawrence (1994), Watt fungi. Actizeta and Chaerodes live in sandy beaches, (1974b). recalling the many species overseas known mainly from
— — arid and semi-arid habitats. Some species (in Art t n t slightly depressed medially; l t r l r n n- dh l p rt t n and C r d l feed on lichens d t n t El tr th n l r h ld r and with striae at night. r l lives in dead tissue of tree ferns and dead weakly defined except for lateral ones. r x l b flower stalks of speargrass. A r Chr p pl th r v t l d b h nd; pr x l pr br d; and tr occur under loose bark of standing dead b th pr x nd x d l p r t d r l trees. G n ph l M p O d d and nt pl Abd n th 5 v ntr t th b l Adeliini are frequently found under stones. n h l nn t and l h occur on flowers. M p occasionally feeds on live plant tissue. SYNOPSIS. Only a single species of prostomid, r - The adventive cosmopolitan elements in our fauna are r h tt is recorded from New Zealand. in the genera Alph t b Gn th r n br and Adults and larvae occur in logs of rimu ( r d r b l of which a few species infest stored products. pr n totara ( d rp t t r and hinau Collecting: hand picking from under stones or logs (at (El rp d nt t which have reached the red stage night with a headlamp); processing organic litter through of decay, breaking up into rectangular pieces with mud- Berlese funnels; pitfall trapping; and sifting sand. like substrate in between, on which they feed.
RANGE. North ®, South ®, Stewart ® RANGE. North ®, South ®, Stewart O Kermadecs ®, Three Kings ®, Chathams ®, Snares ® Kermadecs O Three Kings O Chathams O Snares O Aucklands ®, Campbell ®, Antipodes ®, Bounties ® Aucklands O Campbell O Antipodes O Bounties O
EXAMPLES. AG II AE — Ch r d tr h l d EXAMPLE. r r h tt Pascoe, 1868 (Fig. 222). White, 1846 (Fig. 219); IME II AE — A t z t lb t Pascoe, 1875 (Fig. 220); E E IO I AE — M p REMARKS. In earlier classifications prostomids were l n t (Brême, 1842) (Fig. 221). sometimes treated as a subfamily of Cucujidae [46], prob- ably owing to their strongly depressed body. The genus REMARKS. Keys to subfamilies of Tenebrionidae of the r r is also known from Australia. world (based on adults and larvae) are provided by Watt Suggested colloquial name: red log beetles. (1974a). Keys to New Zealand subfamilies, genera, and species of Tenebrionidae (based on adults) are provided by SELECTED REFERENCES. Pascoe (1868), Schawaller Watt (1992). Doyen t l (1990) discussed the subfamily (1993). classification of Tenebrionidae, with Zolodininae treated as a subfamily (in agreement with Watt 1974a). For cos- mopolitan pest species see Booth t l (1990). [7 ] l OE EME I AE Adults and larvae of Tenebrionidae may be reared on a Fig. 223 diet consisting of wholemeal flour and dried yeast, with Length 6-20 mm Tarsal formula 5-5-4 access to water. Suggested colloquial name: darkling beetles. DIAGNOSIS. d ft l n t bp r ll l f n l nd n f r l p b nt testaceous or fuscous, some- REFERENCES. Doyen t l ( 0 Skopin (1964), times with dark brown pattern, or uniformly black. d Watt (1967, 1968, 1974a, 1988, 1989b, 1992), Watt & ll d r t l d fl x d l n t b f r l Triplehorn (1991). n dth t pr n t M x ll r p lp h n x l d rph n l t lI p th p l nt xp nd d nd tr n t n l Ant nn l n - [ 9] l OS OMI AE ll f l f r 11- nt d E v r bl n z nt r Fig. 222 r l htl r n t near antennal insertions. r - th r x b l ndr l usually longer than broad, widest Length 6-8 mm Tarsal formula 4 4 4 anteriorly or at middle, th t l t r l r n d t n t- DIAGNOSIS. d n rr l l n t bp r ll l l n rr r th n l tr El tr th r l n t th tr n l d pr d l br l reddish brown, f ll d v l p d h nd n r h rt n d nd xp n yellowish brown, or brown, with surface punctate. d bd n th r d d h nd n rounded apically, pr n th l r l htl br d r th n pr n t t with indistinct epipleura restricted to basal half; elytral b tr n l pr d d p t r l t r ll b h nd surface sometimes costate. r x pr j t n th r th d t n t n rr n ; fr nt l p l t r v t p n l nd r nd l n th h nd f r pr nt Ant nn 11- nt d th 3- nt d ll ll n r th p n lt t nt l n t l b; p nl r d th nt b d- b l b d; claws simple or toothed. Abd n th 5 h p d E ll round, projecting. r n t l n- v ntr t nd th n t l ll xp d n l
- 1 — SYNOPSIS. There are 21 species of oedemerid in New elytral width, subquadrate or subcylindrical, with lateral Zealand, of which 18 are native and 3 adventive (Hudson margin smooth. Elytra flat to slightly convex, with larger 1975, Watt 1982a). They belong in 6 genera representing punctures bearing shorter decumbent setae and smaller 2 subfamilies, ACE I AE and OE EME I AE All our punctures bearing longer, secondary elytral setae; epi- native species are oedemerines, and some are restricted to pleura distinct, narrow, extending almost to apex of the alpine zone of the South Island. Of the adventive elytra. Legs long, slender. Procoxae conical, projecting, species, 2 are Australian and the other, our only nacerdine, their cavities open posteriorly and internally; meso- is cosmopolitan. Kuschel (1990) recorded 2 adventive coxae narrowly separated. Tarsi with penultimate and species collected at light from Lynfield, Auckland. antepenultimate segments lobed beneath and with dense, Adult oedemerids occur in coastal habitats, under fine ventral setae. Abdomen with 5 movable ventrites. driftwood just above high tide, and in inland habitats on vegetation; they may be common on flowers or herbage or SYNOPSIS. According to Watt (1987) and Pollock under driftwood. Some species are restricted in habitat, (1995) there are 7 native species of pyrochroid described whereas others range from coastal to inland areas (Hudson and 3 or so undescribed. They belong to 3 endemic New 1975). Larvae live in decaying and/or wet wood, some- Zealand genera of subfamily I I A I AE times in timber partly or intermittently submerged in saline Adults are found under bark of dead trunks and or fresh water, and are common in driftwood along coastal branches, or on foliage or flowers. Watt (1987) found very shores; others occur in decaying Olearia and Dracophyllum little pollen in gut contents, which usually contained logs and dead branches inland (Hudson 1975). fungal hyphae and spores and much amorphous vegetable matter. Larvae are almost invariably subcortical on logs, RANGE. North O, South ® Stewart dead branches, or standing dead trunks, where they feed on Kermadecs ®, Three Kings O, Chathams ®, Snares O decaying cambial matter and fungal hyphae. Larvae of Aucklands O, Campbell O, Antipodes O, Bounties O various sizes are found at most times of the year, indicating that the life cycle may take 2 years or more to complete EXAMPLE. OE EME I AE - Thelyphassa lineata (Fab- (Watt 1987). ricius, 1792) (Fig. 223). RANGE. North ® South ®, Stewart O REMARKS. Haemolymph of oedemerids contains the Kermadecs O, Three Kings O, Chathams O, Snares O irritant cantharidin, which may cause blistering of the skin, Aucklands O, Campbell O, Antipodes O, Bounties O so caution is necessary in handling them. Suggested colloquial name: lax beetles. EXAMPLES. I I A I AE - Techmessa concolor Bates, 1874 (Fig. 224), Exocalopus pectinatus Broun, 1893 (Fig. SELECTED REFERENCES. Arnett (1950), Hudson 225). (1975), Rozen (1960). REMARKS. The New Zealand Pyrochroidae as defined by Pollock (1995) are equivalent to the subfamily Pili- [71] Family PYROCHROIDAE palpinae of Pythidae proposed by Watt (1987). These Fig. 224, 225 authors discuss the relationships between genera and the status of the family, and provide keys to species (adults Length 5-10 mm Tarsal formula 5-5-4 and larvae) and genera. We have followed the family DIAGNOSIS. Body soft, flattened dorsoventrally, concept proposed by Pollock (1995), who studied the elongate, subparallel, with elytra often slightly wid- classification, phylogeny, and geographic history of pili- ened posteriorly from midlength, slightly glossy or matt, palpine genera worldwide, and extended the family limits variously punctate, with decumbent and erect hairs; of Pyrochroidae. coloration uniformly brown or black, or bicoloured with Suggested colloquial name: cardinal beetles. pronotum orange and remainder of body dark. Head ent- irely visible from above, gradually narrowed behind SELECTED REFERENCES. Pollock (1994, 1995), Watt eyes (never abruptly narrowed posteriorly as in Scrap- (1987). tiidae, Fig. 233); clypeus with an anterior membranous area; frontoclypeal suture slightly depressed to dis- tinctly carinate; antennal insertions exposed. Maxillary [72] Family SALPINGIDAE palps with terminal segment expanded and truncate (incl. INOPEPLINAE) apically. Antennae usually sexually dimorphic (males Fig. 226, 227 with increased pectination), filiform, serrate, or pec- Length 2-10 mm Tarsal formula 5-5-4 tinate. Eyes large, projecting, entire or slightly emar- ginate (Techmessodes). Pronotum small, two-thirds of Members of this family are represented by two distinct
—62— forms: smalI carabid-like species, moderately convex to [73] Family ANTHICIDAE slightly flattened, with a distinct `waist' between pro- Fig. 228-231 thorax and elytra, and elytra narrowly oval (Aegialitinae and Salpinginae); and a staphylinid-like species of Ino- Length 2-7 mm Tarsal formula 5-5-4 peplinae with short elytra, exposed abdomen, and strongly DIAGNOSIS. Body antlike, elongate, convex, with flattened body. elytra much broader than head and pronotum, vari- ously punctate, clothed with decumbent and erect hairs; DIAGNOSIS. Body usually coarsely punctate, sub- coloration black, brown, rust brown, or bicoloured with glabrous or with long, protruding setae ( r h - dark or yellowish spots on elytra which sometimes form lp n uniformly black, dark brown, reddish brown, broad bands. Head as broad as pronotum or slightly or bicoloured reddish brown with brown elytral spots. broader, abruptly constricted basally and forming a Head prognathous, never abruptly narrowed behind narrow neck (exception: r d Lagrioidinae); ant- eyes; antennal insertions exposed. Maxillary palps ennal insertions exposed. Maxillary palps with last with terminal segment enlarged, broad and truncate segment usually securiform. Antennae 11-segmented, apically or narrowly elongate (Inopeplinae). Antennae filiform, usually incrassate apically but without a dis- 11-segmented, filiform, without a club (Inopeplinae) or tinct club (a weak 3-segmented club present in M r tr with a loosely formed 3-, 4-, or indistinctly 5-segmented and r d Eyes small to large, slightly protruding club ( r h lp n Pronotum narrowed at base, laterally. Pronotum approximately oval or quadrate in broadened apically. Prosternum long in front of coxae. outline, widest at front and constricted (slightly so in Procoxae globose, narrowly separated, open behind; r d at or near base (C t lateral pronotal mesocoxae globose, and metacoxae transverse; coxae carinae absent. Elytra elongate oblong to elongate or narrowly or moderately broadly separated (Aegialitinae). subparallel (some Anth much broader than prono- Abdomen with 5 ventrites, the 2 basal ones connate. tum or head; punctation confused; epipleura incomplete. Procoxae projecting, their cavities confluent, open behind SYNOPSIS. There are 22 native nominal species of (but closed internally); metacoxae almost touching (Mac- salpingid in New Zealand (Watt l982a); 13 of them are ratriinae) or separated (Anthicinae, Lagrioidinae). Tarsi represented in NZAC. The species belong to 4 genera in 3 with penultimate segment lobed. Abdomen with 5 subfamilies — AEGIALITINAE, INOPEPLINAE, and SALPINGINAE. ventrites, the basalmost 2 connate in r d Kuschel (1990) listed 8 species of S lp n from Lyn- field, Auckland. SYNOPSIS. In New Zealand there are 26 species of Species of S lp n are found on foliage of various anthicid, 9 of them adventive, in 7 genera and 4 sub- trees, shrubs, and herbaceous plants, with some in sooty families — ANTHICINAE, LAGRIOIDINAE, LEMODINAE, and moulds, and are easily collected by beating or sweeping. MACRATRIINAE (Werner & Chandler 1995). Two genera Some larvae of S lp n live under the bark of dead are endemic to New Zealand. Kuschel (1990) recorded 12 twigs. Species of r pn d live under bark of dead species in 4 genera from Lynfield, Auckland. trees, including th f The most specialised species Anthicids are general scavengers, and occur in decaying are in Ant r t d living under stones in the inter- vegetation such as compost or heaps of prunings, on tidal zone on many offshore islands (Watt 1962, 1982b). vegetation near streams, in forest litter, and under logs. They have an elongate terminal tarsomere with strong They may be collected by sweeping vegetation, sifting claws, as in species of Dryopidae and Elmidae [21, 22] — organic matter, or from pitfall traps. r d br n an adaptation for life on rocks in the intertidal zone. which occurs on beaches of the North Island, has related species in Australia and Chile (Lawrence & Britton 1994). RANGE. North ®, South ®, Stewart ® Kermadecs ® Three Kings ®, Chathams ®, Snares RANGE. North ® South ®, Stewart ® Aucklands ®, Campbell ®, Antipodes ®, Bounties ® Kermadecs O, Three Kings ®, Chathams ®, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O EXAMPLES. SALPINGINAE - S lp n b l n t Pascoe, 1876 (Fig. 226); INOPEPLINAE - r pn d f ld EXAMPLES. ANTHICINAE - Anth h p r King, 1869 Waterhouse, 1876 (Fig. 227). (Fig. 228); LAGRIOIDINAE - r d br n Pascoe, 1876 (Fig. 229); LEMODINAE - C t r p (Broun, 1880) (Fig. REMARKS. Salpingidae are poorly known, and the 230); MACRATRIINAE - M r tr x l Pascoe, 1877 (Fig. family is in need of taxonomic revision. 231). Suggested colloquial name: bark mould beetles. REMARKS. New Zealand's anthicids have been revised SELECTED REFERENCES. Spilman (1954, 1967), and described by Werner & Chandler (1995). Young (1991a, b). Suggested colloquial name: ant beetles. - 3- SELECTED REFERENCES. Abdullah (1969), Kitayama SELECTED REFERENCES. Hayashi (1972), Young (1982), Werner & Chandler (1995), Young (1991c). (1991d).
[7 ] l A E I AE [75] l SC A II AE (= EUGLENIDAE = XYLOPHILIDAE) Fig. 233
Fig. 232 Length 1-3 mm Tarsal formula 5-5-4 Length 1-3 mm Tarsal formula 5-5-4 DIAGNOSIS. d ft l htl fl tt n d bl n t IAG OSIS d ft l n t brown or black, often n rr l l n t black, brown, or yellowish brown, with bicoloured elytra, with h d nd pr n t r d- sometimes with dark spots on elytra, uniformly pubescent. l n rr r th n l tr convex, punctate, and th d tr n l d fl x d br ptl n tr t d b h nd r t nd/ r d b nt h r d d fl x d br ptl nt n rr n ; fr nt l p l t r d t n t; n tr t d b h nd M x ll r p lp th l t nt nn l n rt n xp d M x ll r p lp th nt r f r Ant nn 11- nt d f l f r t r n l nt r f r Ant nn f l f r E l r r l f t d r n t ll b d- th t l b E r n t C- h p d r l r t th t l t r l r n El tr br d v l th f t d r n t tr n v r th l t r l r n d t n t h ld r b t l n p pl r ; p n t t n n pl t nt r rl ; disc sometimes with 2 small basal nf d r x pr j t n th r v t p n; impressions. El tr br d pr n t t b with metacoxae separate. r th p n lt t nt confused punctation; p pl r n pl t r x r d d nd th pr d n n l b d Abd n th pr j t n th r v t p n b h nd r th b l t rn t nn t r l dl f d p n lt t nt l b d b n th b l p r ll d v l p d (Fig. 85), p b nt n t rr t Abd n l SYNOPSIS. Watt (1982a) reported 9 native species of v ntr t fr aderid for New Zealand. Fifteen species (many undes- cribed) are represented in NZAC, almost all of them SYNOPSIS. There are 4 nominal species of scraptiid belonging to a genus at present recognised as `Xylophilus.' recorded from New Zealand (Watt 1987), plus several The assignment of New Zealand species to this European undescribed in NZAC. The described species belong to 2 genus is, however, questionable. Scraptogetus (Metasclera) genera in subfamily SCRAPTIINAE. Kuschel (1990) reported is also recorded from New Zealand (Hudson 1934), but its 3 species of Nothotelus from Lynfield, Auckland. assignment to Aderidae is only partially confirmed (Watt Adults occur on foliage and flowers in mainly forested 1987). Kuschel (1990) recorded 5 species of `Xylophilus' areas, and larvae are known from leaf litter.Young (1991e) from Lynfield, Auckland. recorded scraptiid larvae from beneath bark, among Adult aderids are commonly found on foliage, and decaying woody fibres of dead logs, and from lichens. In larvae live in rotten wood, under bark, and in leaf litter. Australia, larvae occur under bark, in rotten wood, or in Adults may be collected by beating foliage, in Malaise leaf litter (Lawrence & Britton 1994). traps, or by light trapping. The larvae are probably asso- ciated with decaying wood (Young 1991d). RANGE. North ®, South ®, Stewart RANGE. North ®, South ®, Stewart ® Kermadecs O Three Kings O Chathams ®, Snares O Kermadecs O Three Kings ®, Chathams ®, Snares O Aucklands O Campbell O Antipodes O Bounties O Aucklands O, Campbell O, Antipodes O, Bounties O EXAMPLE. SCRAPTIINAE - Nothotelus usitatus (Broun, EXAMPLE. `Xylophilus' nitidus (Broun, 1893) (Fig. 1880) (Fig. 233). 232). REMARKS. Scraptiidae may be readily distinguished REMARKS. Aderids are similar in appearance to Anthi- from Anthicidae [73] and Aderidae [74] by the pronotum cidae [73], but are distinguished by having the penultimate having lateral carinae and being equal in width to the tarsal segment vestigial (cf. well developed and lobed) and elytral base, and from Melandryidae [62] by having the the preceding segment lobed, larger and coarsely faceted eyes deeply emarginate, head strongly constricted post- eyes, no elytral epipleura (cf. present but incomplete), and eriorly, and tibial spurs not serrate (Fig. 85). first 2 ventrites connate or solidly fused. This group is very Suggested colloquial name: soft leaping beetles. poorly known in New Zealand. Suggested colloquial names: puppet beetles, ant-like SELECTED REFERENCES. Franciscolo (1964), Watt leaf beetles. (1987), Young (1991e).
- - Superfamily CHRYSOMELOIDEA (PHYTOPHAGA) alus tristis. The majority of our species are cerambycines, Adults are characterised by having pseudotetramerous of which 6 or so species of Zorion are perhaps the most tarsi (5-5-5-segmented with the 4th segment reduced), the strikingly colourful beetles in our fauna. The species of head usually not prolonged into a rostrum, antennae Lamiinae were treated by Breuning (1962), who also filiform, not clubbed, and male genitalia of a reduced provided keys for their identification. cucujoid type (Lawrence & Britton 1994). Larvae lack a Adults feed on foliage, flowers, or bark, and some adults distinct mandibular mola, gular area, and hypopharyngeal feed in live shoots before sexual maturation; larvae feed bracon. All species are phytophagous. internally on bark or woody material of trees and shrubs, A few larvae of exotic species are known to develop in seed pods, or in stems of herbaceous plants (Booth et al. 1990, [76] Family CERAMBYCIDAE Lawrence & Britton 1994). Many species, particularly adventive ones, are of economic importance as pests of [assisted by G. Kuschel] commercial timber. Fig. 234-237 Collected by netting, sweeping foliage and flowers, at
Length 2.5-50 mm Tarsal formula 5-5-5, light, or by rearing larvae from wood. appearing 4 4 4 RANGE. North ®, South ®, Stewart DIAGNOSIS. Body elongate, subcylindrical or flattened Kermadecs ®, Three Kings ®, Chathams ® Snares O and wide, often broad-shouldered, pubescent or sub- Aucklands O, Campbell O, Antipodes ®, Bounties O glabrous; coloration variable through brown, black, or green, or of complex pattern. Head variable in form, EXAMPLES. IO I AE - Prionoplus reticularis White, from prognathous with strongly projecting mandibles 1 (Fig. 3 ; CE AM YCI AE - Zorion sp. (Fig. to hypognathous or slightly opisthognathous; antennal 35 Oemona hirta ( br 1775 (Fig. 3 ; ΑΜΙI ΑΕ insertions on 2 pronounced tubercles or swellings on – l t l costipennis (Breuning, 19 (Fig. 37 frons, either between eyes and mandibles (Prioninae, Spondylidinae) or between eyes (Cerambycinae, Lami- inae). Maxillary palps with last segment tapering apically REMARKS. Important references for identification are or truncate (Cerambycinae). Antennae capable of being Duffy (1953) for British species and imported timber flexed backwards against body, elongate, extending pests, and Duffy (1963, 1968) for Australian and Oriental beyond middle of elytra, often equal to body length or larvae. For world references on economically important much longer. Eyes usually emarginate. Pronotum cerambycids, see Booth et al. (1990). The subfamily clas- sification was discussed by Napp (1994) and Lawrence & cylindrical, with lateral margin distinct, at least basally (Prioninae), to completely absent. Mesonotum usually Newton (1995). with stridulatory files. Elytra variable, from narrow and Suggested colloquial name: longhorn beetles. subcylindrical to broad and flattened (Prioninae), vari- ously sculptured. Procoxae from globular to transverse, KEY TO SUBFAMILIES OF CERAMBYCIDAE their cavities open or closed. Tarsi with penultimate 1 Antennal insertions distant from base of mandibles in segment reduced. Abdomen usually with 5 ventrites. lateral view ... 2 Aedeagal apodemes free, or largly so; internal sac long, —Antennal insertions close to base of mandibles (Fig. with pigmentation confined to struts (paraprocts); styli 234), nearly in contact in lateral view ... 3 usually large. Larvae with ambulatory ampullae present on abdominal segments 1-6 or 7. 2(1) Protibiae not grooved before apex; head weakly pro- gnathous; maxillary palps with apical segment slightly SYNOPSIS. One of the largest beetle families in New expanded and truncate at apex; antennae moderately to Zealand. Hudson (1934) recorded 250 native and 6 ad- extremely elongate ... (Fig. 235, 236) .. CE AM YCI AE ventive species of cerambycid, whereas Watt (1982a) —Protibiae obliquely grooved on internal face before estimated 180 valid nominal native species and 8 adven- apex; head usually hypognathous or opisthognathous; tive. Approximately 196 native and 8 adventive species maxillary palps with apical segment fusiform; anten- are represented in NZAC, belonging to some 56 genera (8 nae usually moderately elongate of them exotic) in 4 subfamilies – IO I AE S O Y - ... (Fig. 237) .. AMII AE I I AE CE AM YCI AE and AMII AE Kuschel (1990) recorded 56 species in 33 genera from Lynfield, Auckland. 3(1 Pronotum with lateral carinae, at least in basal half, The Prioninae are represented by an endemic species, and bearing spines; procoxae strongly transverse; lab- Prionoplus reticularis, which is one of the largest beetles rum firmly united to clypeus ... (Fig. 234) .. IO I AE in our fauna, reaching 50 mm in length. The Spondylidinae —Pronotum lacking lateral carinae; procoxae rounded; are represented by an adventive Palearctic species, Arh p- labrum free ... S O Y Ι ΙΝΑΕ
- 5- SELECTED REFERENCES. Bain (1976a, b), Blair broad beak, notched eyes, enlarged hind femora usually (1937), Breuning (1962), Dumbleton (1957), Hosking bearing teeth, and shortened elytra exposing the pygidium. (1978a–e, h), Napp (1994), Watt (1983c), Zondag & Bain There are 25 native species in 4 genera of Eumolpinae in (1976). our fauna. For a key to genera and descriptions of some species see Shaw (1957a, b). Our 4-9 species in 2 genera of Cryptocephalinae are [77] Family CHRYSOMELIDAE distinct in their partially glabrous, compact, subcylindrical (including UC I AE or cylindrical body form, hypognathous head deeply in- [assisted by L. LeSage] serted into the prothorax, broadly separated long antennae, and often exposed pygidium. Fig. 238-243 Our Galerucinae include some 76 species in 2 tribes, Length 1-10 mm Tarsal formula 5-5-5, appearing 4-4-4 Galerucini and Alticini (the latter is often given subfamily status). Galerucini, represented by some 57 species in 3 DIAGNOSIS. Body form very diverse, elongate and endemic genera, are characterised by their prognathous or subparallel, oval, hemispherical, cylindrical or sub- hypognathous head, with usually filiform antennae inser- cylindrical, flattened or strongly convex, usually sub- ted close together on the frons, pronotum with distinct glabrous but sometimes pubescent or bearing scales lateral margins, and metafemora swollen not much more (Bruchinae, some Eumolpinae), glossy or dull (e.g., some than the mesofemora. Galerucinae); coloration brown, black, yellowish, or mul- The Alticini, commonly called flea-beetles because of ticolored with spots or stripes, often with a metallic shine their usually small size (1-5 mm) and ability to jump, are (e.g., Chrysomelinae). Head prognathous, hypognathous, represented by some 19 species (6 exotic, 13 native) in 10 or opisthognathous, sometimes deeply inserted into genera. They vary in form from elongate to oval and prothorax, sometimes bearing grooves or prominences, convex, and usually are glabrous and variable in colour— sometimes rostrate but with rostrum no longer than wide metallic, brownish, yellowish, black, blue, or dark with (Bruchinae); antennal insertions either close together pale stripes. They are distinct in having usually filiform on from or welI separated, never on a prominence. antennae with narrowly separated insertions, pronotum Antennae not capable of being flexed backwards with distinct lateral margins, and strongly enlarged meta- against the body, usually shorter than body, filiform, femora containing the endoskeletal jumping organ. sometimes broadening apically. Eyes usually entire. Our Chrysomelinae comprise some 41 species (includ- Pronotum narrower than elytra or as broad as elytral base, ing 5 exotic) in 9 genera. They are usually of medium size, with or without lateral margins. Mesonotum lacking predominantly glossy and metallic, with body shape var- stridulatory files. Elytra entire or shortened and leaving iable from narrowly elongate to broad or hemispherical pygidium exposed, striate or with confused punctures .Tarsi and of differing convexity, and head usually oblique, with with 4th segment usually short and concealed at base of front margin of clypeus approximately truncate, antennae preceding segment. Abdomen with 5 ventrites; basal filiform and widely separated, and procoxae transverse. ventrites not fused. Aedeagal apodemes fused to form a The majority of chrysomelids feed on leaves, though spoon-like roof over internal sac, which is short and in some are known to consume pollen and anthers. Bruchines dorsal view completely covered by fused apodemes; feed mainly on legume seeds, and all but one are con- ovipositor with proximal hemisternites short, weakly sidered to be agricultural pests. In New Zealand chryso- pigmented or membranous; styli usually very small or melid adults are commonly collected from low-growing absent. Larvae without ambulatory ampullae on abdom- plants or bushes. Many species are host-specific. The fam- inal segments. ily contains a relatively large number of pests of cultivated plants, some of them transmitting plant viruses. Paropsis SYNOPSIS. Hudson (1934) recorded 155 native species charybdis (Chrysomelinae), self-introduced to New Zea- of chrysomelid and 1 adventive species of bruchid (here as land from Australia, is causing severe damage in gum Bruchinae). Watt (1982a) recorded 146 native species and plantations (Selman 1963, Bain 1977a). For cosmopolitan 7 adventive or introduced, including 1 bruchine. Five species of economic importance see Booth et al. (1990). more introduced bruchines are now represented in NZAC, Larvae have very diverse feeding habits, foraging exter- making a total of some 134 native species and 19 adventive nally or internally on leaves, or stems, or roots. In general, or introduced. These belong to some 33 genera in 5 sub- larvae of Cryptocephalinae feed on dead leaves accumu- families – UC I AE C Y OCE A I AE EUMO I AE lated on the ground (known only from exotic species), C YSOME I AE and GA E UCI AE Kuschel (1990) repor- those of Eumolpinae occur in the soil and probably are ted 20 species in 12 genera from Lynfield, Auckland. root feeders, those of Chrysomelinae feed on foliage, and The Bruchinae, represented by 6 species in 4 genera, are many Galerucinae are miners of roots and leaves, though distinct in having an ovate body, usually broad posteriorly, some feed externally on foliage, fruits, and flowers; all our the head concealed from above and prolonged into a short, introduced Bruchinae feed on legume seeds.
- - Several beneficial leaf beetles have been deliberately —Body oval to rounded or robust, but not cylindrical introduced to New Zealand for weed control: Alticini — (Fig. 239, 243), often strongly convex and constricted A l h r ph l originally from South America, for anteriorly; antennae usually short 5 control of alligator weed (Alt rn nth r ph l x r d and n t r j b from Europe, for control of 5(4) Head oblique, with frontal margin straight or broadly ragwort (S n j b Chrysomelinae — Chr l n arcuate; clypeus distinct, more or less membranous; h p r and C dr n from Europe, for control of body usually broadly oval, robust, slightly depressed, St John's wort ( p r Bruchinae — r h d v l- glabrous; pygidium usually not exposed; 3rd tarsal l from the United Kingdom, for biological control of segment usually bilobed ... (Fig. 239) .. C YSOME I AE broom (C t p r (see Syrett t l 1996). —Head hypognathous, with frontal margin usually broadly emarginate; clypeus indistinct; body elongate- RANGE. North ®, South ®, Stewart ® robust to oval and convex, sometimes pubescent, with Kermadecs ® hr K n ® Ch th ® Sn r Ο elytra often broader than pronotum; pygidium usually Aucklands O, Campbell O, Antipodes O, Bounties O at least partly exposed; 3rd tarsal segment distinctly bilobed ... (Fig. 243) .. EUMO I AE EXAMPLES. UC I AE— r h d v ll (Fabricius, 1792) (Fig. 238); C YSOME I AE - Chr l n h p r SELECTED REFERENCES. Bain (1977a), Borowiec Forster, 1771 (Fig. 239); GA E UCI AE —A l h r - (1987), Jolivet & Cox (1996), Kay (1980), Lawson ph l Selman & Vogt, 1971 (Fig. 240), Ad x v l r (1991), Pfaffenberger (1991), Reid (1995), Seeno & (Broun, 1880) (Fig. 241); C Y OCE A I AE - Arn Wilcox (1982), Selman (1963), Shaw (1957a, b). br n Sharp, 1876 (Fig. 242); EUMO I AE - E l p br nn (Fabricius, 1792) (Fig. 243). Superfamily CURCULIONOIDEA REMARKS. New Zealand's chrysomelids are poorly (RHYNCHOPHORA) known and badly in need of taxonomic revision. Many of [assisted by G. Kuschel and B.M. May] our genera which are now considered as native may reveal a broader origin in the light of detailed studies of the entire Adult weevils are distinct in having the anterior part of the Australasian fauna. head extended into a long or short rostrum (Fig. 6, 7), in Suggested colloquial name: leaf beetles (Alticini — flea- which the mouthparts are often strongly modified (fusion beetles). of labrum, fusion of lacinia and galea, usually reduction of mandibles, formation of rigid palps). Additional diagnos- KEY O SU AMI IES Ο C YSOME I AE tic characters are antennae not filiform, usually clubbed, 1 Body ovate, slightly broadening posteriorly, bearing with a long scape sometimes fitting into grooves on either recumbent setae or scales; head produced anteriorly side of rostrum (Fig. 7), body rigid, with closed procoxal and mesocoxal cavities, procoxae globular with concealed into a short, broad rostrum, strongly deflexed, con- trochantins, 2 or more abdominal sternites connate, and stricted behind eyes into a distinct neck tarsi pseudotetramerous (5-5-5-segmented but appearing ... (Fig. 238) .. UC I AE —Body not ovate, lacking setae and scales (exception: 4-4-4-segmented). some Eumolpinae); head not rostrate, usually not Larvae are distinct in their cylindrical body form (Fig. 99), with abdominal segments 2-7 unsclerotised and with strongly deflexed, without a neck, and usually deeply dorsal plicae or folds, the presence of a hypopharyngeal inserted into prothorax bracon (except in some leaf miners), maxillae with lacinia (1 Antennae closely adjacent, inserted between eyes on and galea united to form a mala, and legs absent or ves- frons (Fig. 240) ... 3 tigial (May 1993). —Antennae well separated, inserted above mandibular bases (Fig. 239, 243) ... 4 [78] Family NEMONYCHIDAE 3(2) Metafemur moderately to strongly enlarged, in Alticini adapted for jumping, with an endosclerite, in [assisted by G. Kuschel] Galerucini not so ... (Fig. 240, 241) .. GA E UCI AE Fig. 244
—Metafemur not enlarged, not adapted for jumping, Length 2.0-2.8 mm Tarsal formula 5-5-5, without an endosclerite ... 5 appearing 4-4-4 4(2) Body cylindrical or subcylindrical, with pronotum DIAGNOSIS. Body moderately elongate, narrowly about as wide as elytral base; antennae usually long oval in outline, rust brown to dark brown, with yellowish- and filiform; pygidium usually partly exposed white decumbent pubescence and some scattered erect ... (Fig. 242) .. C Y OCE A I AE and inconspicuous setae. Head abruptly constricted in
— 7— fr nt f f r n n l n t n rr r tr h r r l f r n p tt rn d pr d d h t fl tt n d p ll slightly shorter than pro- nt r rl nt r tr l n p r d l r t r thorax in male and longer in female; nt nn l n rt n v ntr ll nd ll h rt br d nd fl tt n d (not n p l th rd f r tr d t nt fr nd b l r cylindrical), rarely moderately elongate. br d t n t t b t l t t dth f p br fr fr separated by a groove from rest of head. Mandibles ll n l tr n l r th h rt p -l t well developed. Maxillae with long, flexible palps and a t p x nd p r f t f r p n t r d r ll distinct lacinia. Ant nn tr ht n t lb d (ex- Mandibles acute. M x ll r p lp h rt fl x bl l l ception: slightly elbowed in males of Hoherius), in some rt l t d Ant nn tr ht th p h rt ap- males much longer than body length, th r r l proximately as long as 2 following segments combined, d t n t - r 3- nt d l b nd th p r v d and th l 3- nt d l b E r nd l t- nt r b n d f h d r n t ll th r ll r l f t d r n t n t r n t El tr tr n v r b-b l r n j n d t l t r l r n f d t n tl nd r l rl tr t Hind wings well dev- v r n l n th Elytra internally with a supracostal eloped. Procoxae conical, their cavities closed. b flange, and usually with a scutellary striole. Procoxae th p r n b th x r l l th d nt - globose, projecting, their cavities closed behind. b f r pp nd Abd n l v ntr t fr l rt l t d; l n p r Abd n th 5 v ntr t th b l p d n l d b l tr f d t th r; p d xp d b nd l tr
SYNOPSIS. New Zealand has 4 species of nemonychid, SYNOPSIS. There are 58 endemic New Zealand species of which 1 native species is described and 3 are undes- of anthribid, and 3 introduced (2 adventive, 1 commonly cribed (G. Kuschel, pers. comm.); all belong to the sub- intercepted), belonging to 28 genera in 2 subfamilies — family RHINORHYNCHINAE. ANTHRIBINAE and CHORAGINAE. Of the endemic species, 5 Rhinorhynchus rufulus represents an archaic lineage are confined to the Three Kings, 6 have been recorded only associated with Podocarpaceae (Kuschel 1995). Adults from the North I and 7 from the South I., 6 are confined to occur in epiphytes of podocarp forests and occasionally in the Chathams, 1 is found only on Bounty I 1 is shared by forest litter; a few have been collected in Northland from Stewart I and The Snares, and 1 is present on Stewart I., logs of kauri (Agathis australis). Kuschel (1990) recorded The Snares, and the Auckland Is. The remainder are more this species from Lynfield, Auckland, on tanekaha (Phyl- widely distributed on the main islands. The majority of locladus trichomanoides), miro (Prumnopitys ferruginea), endemic species restricted to a particular zoogeographic and kahikatea (Dacrycarpus dacrydioides). Larvae, desc- region are flightless. ribed and illustrated by May (1993), are pollen feeders Most of our anthribids are associated with standing developing in male cones (strobili) of Podocarpaceae. vegetation in natural plant communities. Holloway (1982) established the following groups of habitats for adults: (1) RANGE. North ®, South ®, Stewart exclusively on standing vegetation; (2) on standing vege- Kermadecs O Three Kings O Chathams O Snares O tation or in litter; (3) exclusively in litter; and (4) in or on Aucklands O Campbell O Antipodes O Bounties O littoral lichens or fungi. Adult anthribids feed on fungi (mainly Ascomycetes and Fungi Imperfecti) and lichens EXAMPLE. RHINORHYNCHINAE — Rhinorhynchus rufulus (Holloway 1982, Kuschel 1995). The fungal material con- (Broun, 1880) (Fig. 244). sists of spores, fragments of hyphae, and ruptured fruiting bodies. Four species feed on hyphae and ascospores of REMARKS. Suggested colloquial name: straight-horned sooty moulds, which grow on the honeydew deposited by weevils. some scale insects (Coccoidea). Larvae are mostly endo- phytic in dead and dying branches of trees and shrubs SE EC E E E E CES Kuschel (1983, 1995), May (Holloway 1982). (1993). RANGE. North ®, South ®, Stewart Kermadecs O Three Kings ®, Chathams ®, Snares [79] l A I I AE Aucklands ®, Campbell O Antipodes O Bounties [ t d b Α ll ] 5 EXAMPLES. A I I AE — Sharpius venustus (Broun, 1914) (Fig. 245); C O AGI AE — Dysnocryptus pallidus Length 0.8-7 mm Tarsal formula 5-5-5, Broun, 1893 (Fig. 246). appearing 4-4-4 DIAGNOSIS. d d r t l l n t t v t REMARKS. For keys to the genera and species occurring d r t l t tr n l nv x usually strongly sclero- in New Zealand see Holloway (1982). tised, l th d th bl br n r n h nd/ r h t Suggested colloquial name: fungus weevils.
— — KEY TO SUBFAMILIES OF ANTHRIBIDAE for oviposition and larval development, pupation occur- [Slightly modified from Holloway 1982] ring in situ (May 1993). Adults of Pachyurinus are often beaten from Podocarpaceae, but the host trees of known First and 2nd antennal segments not arched, usually larvae are angiosperms, Nothofagus and Toronia (May symmetrical, but if asymmetrical then antennae very 1993). Aralius wollastoni lives subcortically in dying or slender and reaching back beyond middle of elytra; dead branches of Pseudopanax arboreus and P. lessonii, these 2 segments no more convex on external margin where the thin bark layer has not yet become dry (Kuschel than on internal margin when antennae folded against 1990, May 1993). For general information on natural body; scrobes either lateral or, if dorsolateral, then 1st history see May (1993) and Kuschel (1995). antennal segment pyriform or obliquely truncate at base ... (Fig. 245) .. A I I AE RANGE. North ®, South O, Stewart O —First and 2nd antennal segments arched and asym- Kermadecs O, Three Kings O, Chathams O, Snares O metrical, more convex on external margin than on Aucklands O, Campbell O, Antipodes O, Bounties O internal margin when antennae folded against body; scrobes always dorsal ... (Fig. 246) .. C O AGI AE E AM ES E I AE - Cyrotyphus tridens ( br 1792) ( 247); AG YCY E I AE - Aralius wollastoni SELECTED REFERENCES. Holloway (1971, 1982), (Sh rp 1876) ( 248). Kuschel (1995), May (1993), Zimmerman (1994). REMARKS. Suggested colloquial name: austral weevils.
[80] Family BELIDAE KEY TO SUBFAMILIES OF BELIDAE [assisted by G. Kuschel] OCCURRING IN NEW ZEALAND Fig. 247, 248 [by G. Kuschel] Length 3-13 mm Tarsal formula 5-5-5, appearing Antennae lacking a distinct club (a weak, loosely defined 4-4-4; or 4-4-4, appearing 3-3-3 4-segmented club rarely present); tarsal formula 5-5- 5, but appearing 4-4-4 because of vestigial 4th seg- DIAGNOSIS. Body elongate and subparallel or broad- ment; tarsal segment 1 at least as long as segments 2 ly, irregularly oval, with head and pronotum distinctly narrower than elytra and tapering anteriorly; surface and 3 combined ... (Fig. 247) .. E I AE especially of elytra sometimes tuberculate, with decum- —Antennae with a distinct 2-segmented club; tarsal formula 4 4 4, appearing 3-3-3 because of vestigial bent or erect pubescence or scales. Head either broad, 3rd segment; tarsal segment 1 shorter than segments 2 abruptly constricted behind eyes, and with rostrum and 3 combined ... (Fig. 248) .. AG YCY E I AE short and broad (Aglycyderinae) or not constricted behind eyes and with rostrum elongate and usually SELECTED REFERENCES. Kuschel (1995), May broad (Belinae); antennal insertions usually in basal (1993), Zimmerman (1994). half of rostrum. Labrum not visible. Antennae straight, with a well defined 2-segmented club (Aglycyderinae) or without a club, rarely with a weak and loosely defined [81] Family BRENTIDAE 4-segmented club (Belinae). Pronotum with sides sub- parallel or converging apically, lacking lateral margins. (incl. ΑΡΙΟΝΙΝΑΕ Elytra variable in shape, often subparallel in part, some- [assisted by G. Kuschel] times with protuberances (Belinae) or with coarsely punc- Fig. 249, 250 tate striae and projecting short hairs and scales. Procoxae Length 2-4//18-75 mm Tarsal formula 5-5-5, projecting, their cavities broadly closed behind but some- appearing 4-4-4 times with narrow lateral extensions. Tarsi with 4th or 3rd segment reduced. Abdomen with 5 ventrites; DIAGNOSIS. Body either extremely narrowly elongate, pygidium concealed. with head as long as pronotum and elytra combined or slightly shorter in females (Brentinae), or moderately SYNOPSIS. Belids are represented in New Zealand by 2 elongate and pyriform, with head as long as elytra or subfamilies – E I AE with 10 described native species in shorter (Apioninae), subglabrous or pubescent. Head 2 genera, and AG YCY E I AE with 1 endemic New with rostrum long and narrow; gular suture distinct Zealand species. Kuschel (1990) recorded the latter from and merged into one; antennal insertions at base, at apex, Lynfield, Auckland, and May (1993) from Whangamoa or in middle of rostrum. Labrum not visible. Maxillae and Cuvier Island. and labium reduced; maxillary palps rigid and labial Adults of Cyrotyphus are usually found in association palps reduced. Antennae straight or (Neocyba) slightly with Podocarpaceae, but utilise dead wood of many kinds geniculate, lacking a club (Brentinae) or club present
- 9- (Apioninae). Prothorax without lateral carinae. Elytra DIAGNOSIS. Body highly variable in form, robust, concealing pygidium. Procoxae projecting, their cav- strongly sclerotised, convex, subglabrous or with scales ities closed. Protibiae without a grooming device on and/or bristles. Head with a long or short rostrum. inner face (consisting of dense vestiture in a groove or Labrum absent. Maxillae reduced, with short rigid impression). Tarsi with penultimate segments bilobed. palps; gular sutures fused. Antennae geniculate, with a Abdomen with first 2 visible sternites fused together, long scape and more or less compact club (Fig. 7, 28). much longer than remaining sternites. Procoxae projecting, their cavities closed. Tarsi with penultimate segment minute and concealed at base of SYNOPSIS. New Zealand's brentid fauna comprises 3 lobed 3rd segment (exception: Platypodinae). genera, with 1 very distinctive native species in subfamily E I AE and 5 species (1 exotic, 2 undescribed) in sub- SYNOPSIS. Curculionids are the most diversified group family A IO I AE of beetles, with some 50 000 species in the world fauna The giraffe weevil, Lasiorhynchus barbicornis, occurs (Lyal 1993). They are also the largest beetle family in New on a variety of trees including kauri (Agathis australis), Zealand. Watt (1982a) recorded 1279 native and 42 ad- pigeonwood (Hedycarya arborea), rewarewa (Knightia ventive species. There are 1496 native and 46 adventive excelsa), tawa (Beilschmiedia tawa), and karaka (Coryno- species in 231 genera represented in NZAC, belonging to carpus laevigatus). The female chews a cavity in the bark 6 subfamilies in the new system of classification proposed of dying or suppressed trees and in freshly felled logs for by Kuschel (1995) — AC YCE I AE (49 genera), CU CU- oviposition (Meads 1976, May 1993). The gorse weevil, IO I AE (126), YO O I AE (Rhynchophorinae) (4), Exapion ulicis, was introduced for biocontrol of gorse COSSO I AE (36), SCO Y I AE (14), and A Y O I AE (2). (Ulex europaeus), and Neocyba metrosideros, a native Kuschel (1990) recorded 184 species (24 adventive) in species, occurs on pohutukawa (Metrosideros excelsa) 102 genera and 6 subfamilies from Lynfield, Auckland. (Kuschel 1990). So far as is known, weevils are exclusively phyto- phagous. They occur in a wide range of habitats: some feed RANGE. North ®, South ®, Stewart O on leaves, others live in the soil and feed on roots; some Kermadecs O, Three Kings O, Chathams O, Snares O bore into stems, and others mine between the upper and Aucklands O, Campbell O, Antipodes O, Bounties O lower surface of leaves (May 1993). Larvae are cryptic, and usually feed internally in plant tissue; some feed on EXAMPLES. Β ΕΝΤI ΑΕ — Lasiorhynchus barbicornis flowers and in developing ovaries (May 1987, 1993). (Fabricius, 179 (Fig. 9 ; ΑΡΙΟΝI ΑΕ — Neocyba metro- Females of Brachycerinae oviposit in the soil, and the sideros (Broun, 1 (Fig. 5 larvae feed on roots (May 1993). The curculionines have many host-specific species, with larvae usually feeding REMARKS. Suggested colloquial name: primitive wee- endophytically on living plants (May 1993). Cossonines vils. and some rhynchophorines utilise dead tissue of bark or rotten wood. Adults of Scolytinae bore through bark to KEY TO SUBFAMILIES OF BRENTIDAE construct a nursery gallery, later expanded by the larvae. Body extremely elongate and narrow, length 18-75 mm; Platypodinae, known as pinhole borers, cause damage to head as long as remainder of body, or nearly so; timber by their tunnelling activities and by `sap staining' antennae without a club; tibial spur present; trochan- from secondary fungal infection, making them considerable ters not elongate, hence femora contiguous with coxae pests in the timber industry. Both scolytines and platy- at base ... (Fig. 249) .. E I AE podines feed on ambrosia fungi cultivated in the galleries. —Body moderately elongate and pyriform, length less Several weevil species have been accidentally intro- than 4 mm; head as long as elytra or shorter; antennae d d nt Ν l nd nd f th r n d r d with a club; tibial spur absent; trochanters elongate, t b p t f r lt r f r tr r rd n r l t f hence femora distinctly disjunct from coxae at base dv nt v p K h l (197 ... (Fig. 250) .. ΑΡΙΟΝΙΝΑΕ RANGE. North ®, South ®, Stewart ® SELECTED REFERENCES. Kuschel (1995), May Kermadecs ®, Three Kings ®, Chathams ®, Snares (1987, 1993), Zimmerman (1994). Aucklands ®, Campbell ®, Antipodes ®, Bounties O
EXAMPLES. Β ΑC YCE IΝΑΕ —Anagotus turbotti (Spiller, [82] Family CURCULIONIDAE 19 (Fig. 51 Mandalotus miricollis (Broun, 1917 [assisted by B.M. May and G. Kuschel] (Fig. 5 ; CU CU IO I AE —Andracalles spurcus (Broun, 1 880) (Fig. 53 Myrtonymus zelandicus Kuschel, 199 Fig. 251-260 (Fig. 5 Stephanorhynchus lawsoni Sharp, 1 7 (Fig.
Length 0.7-30 mm Tarsal formula 5-5-5 or 4-4-4 55 ; COSSONINAE — Macroscytalus remotus (Sharp, 1 7
—7 — (Fig. 256), Xenocnema spinipes Wollaston, 1873 (Fig. 4(3) Mandibular socket deep (deeply emarginate); hypo- 257); SCOLYTINAE — Hylastes ater (Paykull, 1800) (Fig. stomal tooth long, usually sharply pointed; pharyngeal 258), Phloeosinus cupressi Hopkins, 1903 (Fig. 259); process long, about as long as a mandible or longer A Y O I AE- Platypus apicalis White, 1846 (Fig. 260). ( ί 5 57 COSSO I AE —Mandibular socket shallow (shallowly emarginate); REMARKS. The most striking New Zealand weevils are hypostomal tooth absent or indistinct; pharyngeal pro- in the brachycerine genus Anagotus, which includes both cess less than half of mandibular length, or absent ... 5 small species and the largest, often called giant weevils. All are flightless and have specialised habitat requirements. 5( Procoxa antemedian; antennal funicle 5-7-segmented; They are considered endangered because they are prone to first 2 visible sternites fused; sternite 2 distinctly lon- predation by the Polynesian rat (kiore, Rattus exulans), ger than sternite 3; tarsi not or slightly longer than and many are surviving on rat-free islands (see Examples tibiae; sternite 9 of male present of Species and Appendix 1). Evidence that they were once ... (Fig. 258, 259) .. SCO Y I AE broadly distributed on the mainland includes fragmentary —Procoxa postmedian; antennal funicle 4-segmented; remains found in cave deposit at Waitomo (approx. 1700 first 2 visible sternites usually free; sternite 2 not lon- years old) and in the Pureora Buried Forest deposits ger than sternite 3; tarsi distinctly longer than tibiae; formed during the Taupo eruption (circa A.D. 130). sternite 9 of male usually absent New Zealand species of Cryptorhynchini (Curculio- ... (Fig. 260) .. Ρ A O Ι AE ninae), originally treated as a subfamily, were revised by Lyal (1993). New Zealand species of Molytini are revised SELECTED REFERENCES. Bain (1976c, 1977b—e), by Craw (in press), under subfamily Molytinae. Barratt (in press), Craw (in press), Hosking (1978g, 1979), Suggested colloquial name: weevils. Kuschel (1964, 1966, 1969, 1970, 1971, 1972, 1987, 1995), Lyal (1993), Marshall (1926, 1937, 1944, 1953), KEY TO SUBFAMILIES OF CURCULIONIDAE May (1966, 1987, 1993), Milligan (1978, 1979b, c), OCCURRING I NEW ZEALAND Zimmerman (1992—[98]), Zondag (1976, 1977). [Modified from Kuschel 1995]
— v — 1 Epistome (dorso-apical part of rostrum, Fig. 6) usually uneven, raised or impressed in relation to surrounding area; mandibles usually multisetose; rostrum usually E E E CES short and thick, not sexually dimorphic; underside of Abdullah, M. 1969: The natural classification of the elytra lacking stridulatory files; sclerolepidia absent; family Anthicidae with some ecological and etholo- ventrite 9 of male with bladal part extensively sclero- gical observations (Coleoptera). Deutsche entomolog- tised, the arms pigmented, broad, usually discontinuous ische Zeitschrift, MN.F. 16: 323-366. with apodeme ... (Fig. 251, 252) .. AC YCE I AE —Epistome even, not raised or impressed in relation to 1973: The systematic position of Cisidae (Hetero- surrounding area; mandibles usually more sparsely mera) including a catalogue of the world and com- setose; rostrum mainly long and slender, usually sex- ments on central European families of Cucujoidea ually dimorphic; underside of elytra frequently with (Coleoptera). Zoologische Beitrag 19: 189-246. stridulatory files; sclerolepidia frequently present; Allison, A.; Samuelson, G.A.: Miller, S.E. 1993: Patterns ventrite 9 of male with bladal part largely mem- of beetle species diversity in New Guinea rain forest as branous, the arms pigmented, usually narrow, fused to revealed by canopy fogging: preliminary findings. apodeme ... 2 Selbyana 14: 16-20. 2(1) Mesotibia and metatibia with distal or ascending Arndt, E. 1993: Phylogenetische Untersuchungen larval- combs ... (Fig. 253-255) .. CU CU IO I AE morphologischer Merkmale der Carabidae (Insecta: —Mesotibia and metatibia lacking distal and ascending Coleoptera). [Phylogenetic investigations into larval combs ... 3 morphological characteristics of the Carabidae.] Stutt- garter Beitrage zur Naturkunde, Serie A (Biologie) 3(2) Prementum invisible in ventral view, inflexed over 488: 1-56. postmentum; antennal funicle with 6 or fewer seg- Arnett, H.R. 1950: Generic names of the beetle family ments, and segment 7 added to a compact club Oedemeridae and their type species. Journal of the YO O I AE (= Rhynchophorinae) Washington Academy of Sciences 40(7): 217-225. —Prementum visible in ventral view; antennal funicle with 7 segments, if apparently fewer then because of 1973: The beetles of the United States. Michigan, fusion of segments U.S.A., American Entomological Institute. 1049 pp.
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—9 — G OSSA Y O EC ICA E MS b d v r t th t t l t f ll th d ff r nt pl nt n l M t f th d f n t n b l h v b n f r l t d th f n nd r r n th r n t p n nt nd th r r f r n t r n r tt n (199 Sn d r (1935 nd l l t n r ll d n n rr r n h l (19 9 b d v r t f b tl fr p rt l r r l t l nd r nd n dl l b n th l f h b t r pr d t n nd d pt t n f r n n nd t n f th n t n h h th t d h n th r r t ll n n r l d n n pl f h h b t th f n nd fl r f d f n d h b t t r r n th r r p f l n t tr n p r nt ll ; th p n bipectinate ( f nt nn p 1 rh bd bl d l p rt n n t th bl d l p rt n f t rn t 9 adventive ( f p p p l t n h v n rr v d nl cambium, n r f r n nd d ff r nt t n ll n th r ntl n n r nd r d n th r thr h d p r l r t nd r t f v l r pl nt thr h n dv rt nt r d l b r t h n n campodeiform ( f l rv t l t n th rl t d th l p l t r r n d r v d fr th r bl n Campodea ( pl r l n t ll p t r r p rt n f th j l t r d t nd n t n f thr l r t d d r v ntr ll fl tt n d ll l n l d p rt th b l tegmen, h h ft n p d f basat pr d th pr n th h d nd ll th t r n l p (phallobase) nd ll t l t r l l b ll d bd n l pr ( 1 parameres; th penis (median lobe), h h ll n l d cantharidin, d f n v h l f nd n th h l ph t th b b th t n; nd th internal sac (endophallus), f d lt M l d nd O d r d bl t r n h n n h h pr r l br n b t ft n b r nt rn l nth h lf-l l t r l xp n n f th h d pr t ll l r t nd l r t d flagetlum ( ft r r n r tt n v r n th b f th f r t nt nn l nt ( n 199 n br n d 11 1 ll p tr ( f p p l t n r p p n t ll capitate ( f nt nn p 1 nd 3 1 7 x l v r ph r ( f sympatric). p I ll l d nt n r-l tr t r ; ft n alluvial ( f l d p t d b th t n f t r r f rr n t n t h d p l ( 1 7 ambrosia, th f n lt v t d b d-b r n S l t n rd (pl rd n th b l d v n f th x ll n nd l t p d n ; r p f ll th p rt f th f n th t C l pt r ( 5 r t nt th b rr nd t n b th b tl r n (pl r n n l v t d r d r l ampulla (pl p ll bl t rl tr t r ; ft n r f rr n r (pl r n pp nd ( n r ll p r d f bd - t l rv l tr t r ambulatory ampullae n l nt 9 b t ft n pp r n t b l n t bd n l antemedian, t t d b f r th ddl ( f postmedian). nt 1 antenna (pl nt nn p r d nt l pp nd b rn n l d t n l t hn f r r p n t x rd n t th r d f h d nd b r n n r r pt r ( l 17- pr b bl n tr l r l t n h p b d n r f 33 l r t nd d ff r n antennal cleaning organ, n x v t n f th pr t b l n d clavate ( f nt nn p 1 nd 3- 9 th b-l t l fr n f r l n n nt nn n l v rn th nt nn r r l d t n tl l bb d; h n C r b d Clavicornia, r p f b tl th th f t r antennomere, n f th nt nn l d v n ( ll 11 n l h n n d lt El t r d nd E n d - C l pt r h h r j nt d t th r nd f r th nt r pl x `p nd t tr t r n t n f th l n pr t rn l nt nn ( l pr nd t rn l v t ; dd n f r n f th pr nt r r p t (= nt r r t nt r l p t ll r nd nt rn l t f th v t n nv l v v nt f th d pr n b t n l p nd fr n ( l t nt r l p t pr th r x r l t v t th h nd b d ft n r lt n n j p ( l nd/ r n d bl l ( 5 p d n n r th f th n t x l t n ( v t h h l b ( f nt nn p 1 nd 3-31 l r tt h d clypeus, p rt f th n t h d p l b l th fr n t apomorphy, d r v d h r t r r h r t r t t ( f plesio- h h th l br tt h d nt r rl ( 1 morphy); h n apomorphic. coeval ( f t r r r n h v n v lv d nt p - approximated, n l ph l r l t n h p r n l n th l l t l apterous, th t n b r r p t h f p l d p n ll n t b articulation, b l j nt b t n t r d tr t r ( 1 aspirator, t n d v f r pt r n ll n t compound eye, n n rthr p d n r t n f v l Barber's fluid, b tl r l x n fl d th n l 95% (53 p rt l nt (ommatidia) h f h h rr p nd th n l t r ( 9 p rt th l t t (19 p rt nd b nz l (7 p rt f t f th rn Berlese funnel, p 17 n d x l p r f f n biforous ( f p r l th t xt nd d p n n r th connate ( f t t v bl j n d t th r f th r n b t n pp r n l r t th d bl p r l l v bl ( 91 nd p r t nf r t n coronal suture, the median unpaired part of the epicranial endophallus, part of the aedeagus. suture. endopleuron (pl., endopleura), the reduced pleuron of Poly- costate, with longitudinal raised ribs or ridges (costae), much phaga which is fused with the trochantin and hidden internally. coarser than carinae. endosclerite, an internal sclerite, e.g., of the metafemur in coxa (pl., coxae), the basal segment of the leg, articulating to Alticini, sometimes called a metafemoral spring, which the body (Fig. 57-65). constitutes a skeletal jumping organ. coxal cavity, the opening or space in which the coxa articu- epicranium, upper part of head capsule from frons so neck, lates. May be laterally closed by sterna alone, i.e., excluding including frons, vertex, and genae (Fig. l, 6, 7). Hence pleural elements, or laterally open, when only partly or epicranial stem, οr r n l t r nd epicranial suture, a completely closed by pleural elements (Fig. 40-45). Y-shaped suture on the head capsule above the antennae (Fig. crenulate (of margins), evenly notched with small, rounded 113). teeth (Fig. 163, 164 — hind margin of pronotum). epigeal, living on the surface of the earth (cf. hypogeal). Cretaceous, third period of the Mesozoic Era, extending about epimeron (pl. epimera), the posterior division of a thoracic 135-70 million years before the present. pleuron, delimited anteriorly by the pleural sulcus (Fig. 2, 3). r pt h dd n r n l d r f rr n t ( l f h b t οr (b epiphyte, a plant growing nonparasitically upon others, taxonomic status, e.g., cryptic species. especially in the upper branches of trees, e.g., some ferns and crypto-, hidden, as in, e.g, cryptopleuron. orchids. cryptonephridism, a condition in which the Malpighian tubes epipleuron (pl., epipleura), in adult beetles, deflexed or are closely associated with the rectum, forming a convoluted nfl x d l t r l p rt n f pr n t οr l tr ( 3 3 layer over its surface. 35); hence epipleural carina, a sharp ridge at lateral edge of cucujoid (of the aedeagus), a configuration in which the epipleuron. phallobase forms a ring around the penis. episternum (pl., episterna), anterior sclerite of pIeuron, cupule (of antennae), transverse and concave segment delimited posteriorly by pleural sulcus (Fig. 2). preceding the club (e.g., in Hydrophilidae, Fig. 27); hence eplstome, sclerite immediately behind labrum (Fig. 6, b). cupuliform. eruciform (of a larva), resembling a caterpillar (Fig. 97). cuticle, outer layer of the insect i ntegument. excavate(d), hollowed out, concave, as of a margin. decumbent, bent downwards. exoskeleton, extemal skeleton of arthropods, consisting of hard deflexed, abruptly bend downwards. or flexible cuticle to the inside of which muscles are attached. denticulate, beset with tiny teeth or notches. explanate (of a margin), spread out and flattened (Fig. 217 — detritivore, an animal that feeds on organic detritus. pronotal margin). dichotomy, a branching of a single stem into two equal and exuviae, the moulted, cast-off skin of larvae or nymphs. diverging parts; hence dichotomous key, a key with couplets femoral lines, fine ridges on basal abdominal sternite running divided into pairs of opposing character sets. backwards and outwards from inner end of metacoxae (Fig. dimorphic, existing in two forms; hence sexually dimorphic, 92). differing between male and female. femur (pl., femora), third and usually stoutest segment of the disc, medial surface of a body sclerite; e.g., central surface of leg, articulated to trochanter proximally and to tibia distally pronotum. (Fig. 57). disjunct, separated or lacking continuity; often referring to filiform (of antennae), see p. 14 and Fig. 17, 18. distribution of species or populations. flabellate (of antennae), see p 14 and Fig. 33. distal, near or towards the free end of any appendage; that part flagellum (pl., flagella), third part of antenna beyond pedicel. of a segment farthest from the body (cf. proximal). Also part of the aedeagus. diurnal, active or habitually flying by day (cf. nocturnal). flight interception trap, see p. 17. dorsal, on the functionaIly upper surface (cf. ventral). Hence fossorial (of legs), adapted for burrowing in soil. dorsoapical, on the upper surface and towards the apex; fovea (pl., foveae), a pit; e.g., on head and pronotum of dorsolateral, at the top and to the side; dorsoventral, in a line Pselaphinae, Staphylinidae (Fig. 126). from the upper to lower surface. frons, upper anterior portion of head capsule, usually a distinct dorsum, the anatomically upper surface. sclerite between epicranium and clypeus (Fig. l). ectoparasite, an external parasite (cf. endoparasite). frontoclypeal suture, suture between frons and ctypeus (Fig. elytron (pl., elytra), the leathery forewing of beetles, serving as l). a covering for the hind wings (Fig. l). Hence elytral suture, a fungivore, fungus-eater; hence fungivorous (= mycophagous). median ridge dividing the elytra symmetrically (Fig. 93, 94). fuscous, dark brown, approaching black. empodium, a pad or extension between the tarsal claws. fusiform, spindle-shaped, i.e., broad at middle and narrowing endemic, pertaining to species or higher taxa confined to a towards the ends. geographical area under discussion; hence endemism. galea, outer lobe of maxilla (Fig. 5).
-9 - gall, abnormal growth of plant tissue caused by various organ- hypogeal, subterranean (cf. epigeal). isms, including insects. hypognathous (of the head), vertical, with mouthparts facing gena (pl., genae), cheek, that part of head capsule on either downwards (Fig. 9, 10). side below eye (Fig. 2, 4). hypomeron (pl., hypomera), epipleural fold, or in adult beetles geniculate (of antennae), see p. 15 and Fig. 6, 28. (Polyphaga) lateral portion of pronotal disc extended ventrally genital segment, modified 9th abdominal segment in beetles, on either side which meets lateral part of sternum (Fig. 3). containing the genitalia; in the male the 10th segment is hypopharynx, a median lobe immediately behind mouth, usually highly reduced or fused to segment 9. bearing taste receptors. Hence hypopharyngeal bracon (of genitalia, structures relating to reproduction. beetle larvae), a transverse brace between hypopharynx and gibbosity, irregular protuberance; hence gibbose, very anterior part of hypostomal margin. crooked, twisted. hypopleurum (pl. hypopleura), in beetle Iarvae, area below glabrous, without setae. ventrolateral suture. globose, nearly spherical. hypostoma (pl., hypostomata), region of subgena behind mandible. Hence hypostomal margin (of beetle larvae), glossa (pl., glossae), tongue; inner pair of lobes at apex of ventral marginal thickening of the epicranial halves; hypo- prementum of labium. stomal tooth (in some groups of Curculionoidea), a tooth Gondwana (`Land of the Gonds'), during much of the situated in lower mouth between mandibles and base of Mesozoic Era, a southern supercontinent consisting of maxillae. continental blocks of South America, Africa, Madagascar, incrassate (of antennae, etc.), thickened, suddenly swollen India, Antarctica, and Australia with New Zealand. towards the apex (Fig. 134). grooming device, area of vestiture in a groove or impression indigenous, originally inhabiting a particular geographical area on inner face of protibia, usually for cleaning the antennae. (but not necessarily restricted to that area). gula, in prognathous insects, fused lower ends of post-occiput, instar, life stage between moults in a nymph or larva, forming a ventral plate (Fig. 2, 4). Hence gular suture, line of numbered to designate developmental sequence; e.g., the first division between gula and genae; ventro-anterior extension of instar is the stage between the egg and the first moult. postoccipital suture. integument, outer layer of an insect, comprising epidermis and haemolymph, watery body fluid in insects corresponding to cuticle (see also exosketeton). blood. internal sac, part of the aedeagus. habitus, general form and appearance. intersegmentaI membrane, flexible, infolded membrane hemisternites (of female genitalia), two sclerites of sternum 9 connecting segments of body and appendages which allows surrounding vulva. freedom of movement (Fig. 91). heterogeneous, differing in kind, unlike in quality. Jurassic, second period of Mesozoic Era, between Triassic and heteromeroid (of trochanterofemoral junction), strongly Cretaceous, extending about 180-35 million years before the . femur and coxa oblique, so that there is direct contact between present. (Fig. 58, 59). labium (pl., labia), lower lip, or fused second maxillae, heteromerous, with the three pairs of tarsi having different forming floor of mouth in mandibulate insects, behind first numbers of tarsomeres, e.g., tarsal formula 5-5-4. maxillae and opposed to labrum. Hence labial palp, a paired, Hexapoda, superclass of phylum Arthropoda, including 1-4-segmented sensory appendage of labium. Collembola, Protura, Diplura, and lnsecta, characterised by labrum, upper lip, joined lo clypeus in front of mouth (Fig. 1). mouthparts consisting of mandibles, maxillae, and labium, lacinia (pl. laciniae), bladelike inner lobe of maxilla, articu- head with anterior and posterior tentorial arms, 3-segmented lated to stipes and bearing brushes of hairs or spines (Fig. 5). thorax, normally with 3 pairs of legs in the adult, and abdomen with 6-I2 segments, with at most rudimentary limbs. lamella (pl. lamellae), a thin, platelike or leaflike process. Hence lamellate (of antennae), with apical segments having hinge mechanism, a structure of the medial loop of the hind flat, leaflike projections on one side which sometimes cannot wing used for wing folding (in Archostemata and Adephaga) (Fig. 30) but usually can (Fig. 31) be brought closely together. (Fig. 51). lateroventral, lateral and below median horizontal plane. Holarctic, faunal region comprising the northern part of the Northern Hemisphere. ligula (pl., ligulae), collective name for glossae plus para- glossae, h th r f d r p r t ( ί holometabolous, passing through a complete metamorphosis in which the larva is very different from the adult and littoral, in freshwater ecosystems, in shallow water where light transforms dramatically by means of a pupal stage. penetration extends to the bottom sediments, which are col- onised by rooted plants; in marine ecosystems, intertidal, humeral angle, anterolateral portion of elytra. inhabiting the shore. humerus (pl. humeri), shoulder, anterior basal portion of wing. locking mechanism, a structure preventing movement, e.g., hydrophilous, water loving, living at least partly in water. pronotal locking mechanism in adult ptilodactylid and hyphae, threadlike strands making up the mycelium of a chelonariid beetles (Fig. 163, I64). fungus.
-93- l l b f th nd bl r j n n ln n l ( f p b r n n b t n t n r l M l tr p p 17 v l d b l l nt t M lp h n t b nt rn l x r t r tr t r r v n n t pl r l t r n Ad ph Ar h t t nd p r t fr th b d fl d M x ph (C l pt r t r p r t n pr n t l p pl r n nd bl f r t p r d thp rt t t nd j l n fr pl r n ( h n n t ( 1 n mandibular socket, n n t t rn l t r n l ph t r b t n h p r n pr n r r n t n nt n n nd b l r b f pr n t nd pr t rn h r th pl t t d r tl x ll (pl x ll nd p r f j n n t th ( n t p r t d b n xt rn l pl r n h - n thp rt ; r pr nt d n ll n t th f n t n l n t (pl n t d r l l r t f th r nt ( 1 — thp rt ( 5 n maxillary palps, p r d 1-5- pr n t nt d ( ll 3 r n C l pt r n r pp nd bl n ll f t r f v n t n n h nd n f Ad ph f th x ll ( - x ll l b l pl x p nd tr t r n t n f p t p t r r p rt f p r n b t n v rt x nd n x ll r nd l b l l nt ( n l rv f n h d r r l pr nt d t n t l r t ( n occipital d f fth l n t d n l v n t (M ll d v d d nt carina, ll d f n d l v t d r t l n h nd r n f n r ll nv x nt r r br n h (MA nd n v p t ( n d d p t r r br n h (M ( 7 ll (pl ll n d lt n t pl n t n f d l n r r t th ddl f tr t r r r n l b dl l n rr n n l r n ll r p ; n d l l p oblongum cell. h l t b l l rv stemma. d n l b p rt f aedeagus. l r nth l b p rt ll r pl t l d v d n h br n th n nd r r l tr n p r nt ( f n ; ( n G r n d n br n d — 11 f th n nd pl bl ( f t l Ol n l l p h b t - 5 ll n r b f r nt (pl nt p rt f l b pr x l t pr nt th pr nt tt h d t nd t f d th b nt ( t d (pl t d v l ` ll r l nt f th - pr f x ft n p rt n n t th ddl nt f th rthr p d p nd th r x d n ` x ` p r n n f r ( f l rv h p d l l t r (Oniscus p M z l l r b t 5-7 ll n r b f r d pr d nd br dl p ndl - h p d; El d th pr nt p r l (pl p r l l d r v r; n l rv l r p d t - pr f x ft n p rt n n t th h nd nt f th v ntr l fl t v r t th h b r nt n n th r tr t d n l th r x n ` t x ` t p r n ll p th n th ( f h d th p t r v ntr l p t n f t rph r d l h n n f r p ll th thp rt r lt n fr d fl x n f th f l r n h n l rv —p p — d lt n holometabolous n t r n p d f nt n b ld n n l l t r lpt r rn r p t xt r l f t r f n t nt nt v p t t l r t v r ll h rt h rl tr t r ; h n p lp r l r t b r n p lp p th p lp-b r n microsetose. tr t r f th nt l r d d r r h n d r nd n rf f nd bl p r l (pl p r l p r d l b l tr t r t p x rr -p nd n t b l f x ll h n th nd bl f pr nt nn t d t th r d t l nd th r p nd fr r t -j nt d; rr p nd n t l f x ll ( n l f r ( f nt nn p 1 nd 19 5 n ph h t- p f ; r tr t d t n p f f d p r ph l t ( f t x n r t x n t n l d n ll th d n- d nt f n n t r n t n t r l r p pl nt r n l ( f polyphagous). n t p ( f t x n nt n n nl n b rd n t p r pr t th h lv f th n nth t r t t x n n f nl n p ( f polytypic). p r t d n nt rn l r xt rn l p r t th t v nt ll ll lt t b r n n r t t h t r n n (pl n n n f v r t f p l p r t r t l t r l r n l r n f n t l t r l l r t f p t- h p d r pt l d t rr b t f n bd n l nt n b r b tl (S l t n p t n t ( f nt nn th n d f h nt ph f n - t n (= f n v r pr tr d n r tl f r n r f l n h rp p nt l th t th f b ( r ph l nt-l v n ; ppl d t n t th t l v n nt n t p d l nd nt f th n t nt nn pp rt n th fl ll ( 1 n r h v n n r d nt l tr l th r f p rl p d n l t l r t l l tr t r ll pp rt n n n h f n t n l p t n f n r n n n t r n r th r tr t r ; h n pedunculate. n t rn l t v t n ht ( f diurnal). ph ll b p rt f th aedeagus.
-9 --9 - ph r n l pr h ll br n tr t r pt r th r x f d th r x nd t th r x pr j t n nt th r l v t fr th nt rn b l n l f p b nt l th d th f n h r th nd bl ; pr nt n n rl ll C r l n d b nt r p n t t th p n t r -I d pr n f th nl r d nt r n th r b tl nt nt phl v l tr n p rt n d lv d n tr nt fr l v t p p n t v tr n f r t v t f h l t b l n t r t n v l r pl nt nt r d t b t n l rv nd d lt; h n pupal cell, phr t nd r r nd ll n r f r n t th t v t n th l οr d f pl nt t r l r d br n h h h b t t b t n th p rt l f r alluvial d nt th l rv p p t ; puparium (pl p p r f n l- n t r l rv l ph l n v l t n r h t r f r p f r n nt nt th n h h p p t n r ph t ph f d n n/ n pl nt ; h n phytophage. p d (pl p d t r f l t v bl bd n l pl d n x t f n t l f pr t pl nt; n n d lt b tl th nt l ft xp d b h r t r t f th r th ph th t r n t n f th l tr ( 1 9 3 pl rph n tr l h r t r t t ( f apomorphy). p p d p r d f tl r n t h nd nd f bd n f pl r t pl r n rt n l rv t n l t n pI r n (pl pl r n d lt f Ad ph v ntr l t r l p rt p r f r p r- h p d f th r ll t n p pl r n I t r ll nd t rn r d l ll ( tr t r f r d r nd l r t d d ll ; p r t d fr pl r n b n t pl r l t r nd p nt d bl d n (pt r t pl n p rt n f ld n fr t rn b pl r t rn l t r ( th n p x b n th th l tr pl (pl pl f ld r r n l r d l l p h nd n tr t r n b tl n t n f th pl ( f nt nn p 1 radial bar, h h nd br ptl n ft r nt r n th ptero- p t r l t r l t rd th r r nd t n d ; l rl th stigma, nd th pt r t t lf tr n f r d nt th radial -d r l - d n -v ntr l t cell ( -51 p t d n t t d b h nd th ddl r pt r l ( f l d pt d f r z n nd h ld n pr ( 11 b p t nt b l p rt f l b pr x l t pr nt r b nt l n d n r l n n p t l r b h nd th p nd r l t rv v n n l t n pr d pr n p n th r r n C r b d t St ph l n d rh bd t f ph t r pt n n th p nd pr nt r n f l b d t l t nt f r d b r p r n l v n n b n f t r r palpigers nd l b l stipites. r tr (pl r tr n tl pr j t n r r d xt n n pr p d p rt f l t bd n l t r p r t d fr f th h d b r n th thp rt t th nd n d lt p d b tr n v r t r v l ( 7 ; h n rostrate. pr - pr f x ft n p rt n n t nt r r nt f th r x p f r t r b l nt f n t nt nn ( 1 n procoxa, proepimeron. l r t n d r t pl t f th b d ll b nd d b - pr n th ( f th h d h r z nt l th thp rt br n r b t r d r t d f r rd ( l r l p d r f p t th p l t r l t th t p t rn l t r n v l pr n t pp r nd d r l p rt f pr th r x ( 1 n pronotal carina, b l tr n v r r n n Anthr b d l r t t n h rd n n f th t l nv lv n d v l p nt ( 5 nd t r f rr n t l t r l r n f f r l n b t n pr t n h n pr n t ( 1 ; pronotal comb, r f p n n p t r r r b r v f r r pt n r n l nt f n r n f pr n t n t l d t l d ( 1 3 ; pp nd ; n d lt C r l n d p r f r v n n pronotal epipleuron, n d lt Ad ph d fl x d r nfl x d th r d f th r tr n h h th nt nn n r t ( p rt n f pr n t ( 7 d pr t rn t rn f th pr th r x ( 3 3 -39 t ll (pl t ll n ll h ld- h p d pl t ; n prosternal process, p t r r pr l n t n f p t r r d v n f pt r th r n t ; n d lt b tl pr t rn b h nd pr x tr n l r p t b f pr n t nd b t n l tr ( pr th r x f r t th r nt b r n nt r r l b t n 1 n scutellary striole. n ( 3 3 -39 scutum (pl t nd d r l l r t f th r x nd pr x l n r r t rd th n h r d nd f n pp nd ; t th r x th ddl d v n f th n t th t p rt f nt l t t th b d ( f distal). r f r h t h t- h p d; th t r n l nt f th p d t tr r ( f th t r h v n pp r ntl n- p lp f b tl ( 15 t t nt rt l lth h 5 r t ll pr nt ( 7- 1 n r (pl n r n r r pt r v bl p d tr r ( f th t r h v n pp r ntl 3 rt l r l r p n n v r d b br n n nt nn r l lth h r t ll pr nt ( - n r t t d pl n r n ft n pt r t p nt d p t n nt r r r n f n nt t r t h r pt r r t -p n t t th p n t r rr n d n striae. -95- serrate (of antennae, etc.), see p. 14 and Fig. 20, 21. tegmen, part of the aedeagus. sessile, attached directly, without any stem or petiole. tentorial pit, lateral depression in cranial wall usually near or seta (pl. setae), a hairlike projection of the cuticle; hence at frontoclypeal suture (Fig. 1). setiferous, bearing a seta or setae; setose, furnished with setae. tergite, see tergum. sinuate (of edges and margins), wavy. tergum (pl., terga), upper or dorsal surface of any body slime moulds, Mxomycetes, one of the four fungal phyla; segment of an insect, consisting of one or more sclerites. hence slime flux, a plasma-like stage of slime moulds. terrestrial, living on land (cf. aquatic); sometimes taken as the sooty moulds, fungi of the Ascomycetes: Capnodiaceae and complementary opposite of marine, i.e., including freshwater. related families. testaceous, brownish yellow. spiracle, external opening of a trachea. thorax, middle portion of insect body between head and abdo- spore, among fungi, a microscopic structure functioning in men, consisting of three segments, each of which usually bears dispersal and vegetative reproduction. a pair of articulated legs (Fig. 2). spring mechanism, in Polyphaga, an elastic structure of the tibia (pl., tibiae), fourth segment of Ieg, between femur and
me dian loop used for folding the hind wing (Fig. 51). tarsus (Fig. 57). Hence tibial spur, a spinelike projection at or spur, a thick cuticular appendage or spine connected to the near end of tibia. body wall by a joint, especially on the tibia. tomentose, covered with short, matted, woolly hairs. stemma (pl., stemmata), a simple eye of the often circular, tooth (in insects), a short, pointed process from an appendage lateral eye-groups in holometabolous insect larvae. οr r n; v r t t p l th bl nt p x sternite, see sternum. trachea (pl., tracheae), a spirally ringed, internal elastic air sternum (pl., sterna), entire ventral division of any body tube in insects; an element of the respiratory system. segment. trilobed (of the aedeagus), consisting of a ventrally sclerotised stipes (pl., stipites), basic sclerite of maxilla, articulating partly phallobase to which a pair of freely articulated parameres are with head, partly with cardo, and bearing galea, lacinia, and attached, and a simple penis lying above it; generally maxillary palp. considered to be the most primitive configuration in beetles. stria (pl., striae), in general, any fine longitudinal impressed trochanter, second segment of insect leg, between coxa and line; in beetles, a longitudinal depressed line or furrow, often femur (Fig. 57-66). punctured, usually extending from the base to the apex of the trochanterofemoral, pertaining to trochanter and femur con- elytra; hence striate. jointly. stridulatory apparatus, structure used in making sounds, trochantin, a small sclerite separated from pleuron by a mem- consisting of a file-like organ and an opposing scraper or rasp. brane and connected by articular condyle with coxa (Fig. 57). striole, rudimentary or reduced stria. trophic, pertaining to food or eating. stroma (pl. stromata), mass of vegetative fungal hyphae truncate, cut off squarely at the tip. bearing fruit bodies and/or spores. tubercle, a small, knoblike or rounded protuberance; hence stylus (pl. styli), a small, pointed, non-articulated process. tuberculate. sub-, prefix denoting under, slightly less than, or not quite. Tulgren funnel, funnel-shaped device for sorting Iiving insects subcortical, beneath the bark of a tree. from organic debris by means of light and heat. submentum, basal part of labium by which it is attached to urogomphi (sing., urogomphus), in larval beetles usually head capsule (Fig. 2). paired posterior processes on tergum of ninth abdominal segment, either jointed and movable by muscles or unjointed sulcus, a groove or suture marking the line of fusion of two and immovable. formerly distinct plates or structures. UV light-trap, see p. 17. suture, a seam or impressed line indicating where two sclerites join. Hence sutural flanges, posterior sutural edges of elytra, venter, entire undersurface of body of an animal; hence separated by the sutural gap (Fig. 93). ventral. synapomorphy, a shared derived character. ventrad, towards the venter. systematics, study of biological classification based on ventrite, sternal sclerite of abdomen; the first ventrite is equi- relationship through inherent similarity; hence systematic valent to the first visible basal sternite. sequence of taxa, as used in this volume. vertex, top of head between eyes, frons, and occiput, anterior tactile, pertaining to sense of touch, or used for touching. to occipital suture. tarsus (pl., tarsi), terminal leg segment attached to apex of vesicle, a small sac or bladder. tibia, consisting of 1-5 articles or tarsomeres. Hence tarsal waist, basal constriction of pronotum in beetles, e.g., formula, the number of tarsomeres on the protarsus, Anthicidae (Fig. 228-231). mesotarsus, and metatarsus. weevil, beetle of the family Curculionidae. taxonomy, theoretical study of classification, including its window trap, see p. 17. bases, principles, procedures, and rules. wireworm, soil-inhabiting larva of Elateridae feeding on roots.
-9 - KEYS O AMI IES O CO EO E A Suborder Polyphaga K OW OM EW EA A 5(1) Antennae with a lamellate, flabellate, or exceptionally pectinate club (Fig. 30-33) ... (p. 98) .. Group A —Antennae not as above .. 6 I: Y AMI A KEY [pr v d d b Kl z ] 6(5) Tarsi 5-5-5-segmented but appearing 4-4-4-seg- 1 First ventrite divided by metacoxae (Fig. 88, 89); mented, with segment 4 reduced and segment 3 prothorax usually with distinct notopleural sutures, distinctly bilobed, not so in some Chrysomelidae distinct from sharp lateral margins (Fig. 2); tarsi 5- (exception: weevil groups Belidae : Aglycyderinae segmented; antennae usually filiform (Fig. 17, 18); and Brentidae : Apioninae (Neocyba), in which tarsi medial loop of hind wing with a hinge mechanism appear 3-3-3-segmented but then head with a rostrum) (Fig. 51) ... (p. 20) ADEPHAGA .. 2 (Fig. 67-81) ... (p. 98) .. Group B —First ventrite usually undivided by metacoxae and —Tarsi not as above, if 5-5-5-segmented with segment 4 continuous from side to side (Fig. 90); prothorax reduced then all segments simple, and if appearing 3- without notopleural sutures — ventral portion of notum 3-3-segmented then head without a rostrum ... 7 (hypomeron) jointed direcly to sternum on either side 7(6) Body bullet-shaped, with sides subparallel and by notosternal suture (Fig. 3); tarsi and antennae often not as above; medial loop of hind wing with a spring abdominal apex triangular; head deflexed; abdomen with first 2 ventrites strongly connate, the suture mechanism (Fig. 51) ... (p. 23) POLYPHAGA .. 5 partially obliterated; metasternum with a distinct transverse suture ... (Fig. 157) .. [19] BUPRESTIDAE —Body not so; first 2 ventrites rarely fused ... 8 Suborder Adephaga Superfamily Caraboidea 8(7) Tarsi usually 5-5-5-segmented, with terminal seg- 2(1) Terrestrial beetles with legs adapted for crawling or ment enlarged and ending in strong claws, often longer running; metacoxae not meeting elytra laterally (Fig. than all other segments combined (Fig. 160); aquatic 2); antennae largely pubescent; dorsal surface with or semi-aquatic beetles broadly oval in outline, pronounced, erect sensory setae (Fig. 107) ... 3 convex, with deflexed head more or less concealed by —Aquatic beetles with at least hind legs adapted for pronotum from above; legs retractable into body swimming, often flattened and with dense pubescence cavities (Fig. 52, 53); prosternum with a process (Fig. 108-110); metacoxae extending laterally to meet fitting into a cavity of mesosternum (Fig. 52, 53); elytra (Fig. 88, 110, b); antennae entirely glabrous, or antennae weakly clubbed (3-7 segments), comblike, nearly so; dorsal surface without pronounced, erect sawtoothed, or distinctly clubbed (Fig. 29) sensory setae (phreatic species, e.g., Kuschelydrus, ... (p. 99) .. Group C with long, fine, setae on sides — Fig. 109) ... 4 —Tarsi other than 5-5-5, or if 5-5-5-segmented then terminal segment not enlarged; sometimes segment 4 reduced and tarsi appearing 4-4-4-segmented (Phalac- 3(2) Metacoxae widely separated; antennae thick, mon- ridae); other features not as in above combination... 9 iliform (Fig. 19); pronotum usually with 3 deep longitudinal grooves, or at least 1 groove; head with a 9(8) Maxillary palps usually longer than antennae (Fig. distinct neck; metasternum without transverse suture 112, 116) (exception: Meropathus, with maxillary ... (Fig. 101) .. [1] RHYSODIDAE palps much shorter than antennae — Fig. 117); an- —Metacoxae contiguous or narrowly separated (Fig. 2); tennae 7-11-segmented with a 2-5-segmented club; antennae slender, filiform (Fig. 1); pronotum without hydrophilous species, spherical or moderately convex deep grooves; head usually without a distinct narrow to ovoid and flattened ... (p. 99) .. Group D neck; metasternum with a transverse suture (Fig. 2) —Maxillary palps shorter than antennae, inconspicuous; ... (Fig. 102-107) .. [2] CARABIDAE antennae usually 11-segmented, with a 3-segmented club or unclubbed; shape usually not as above ... 10 4(2) Antennae longer, filiform; eyes not divided into two; forelegs short; hind legs long, modified for swimming, 10(9) Pronotum with posterolateral angles strongly pro- flattened and fringed with hairs duced behind (Fig. 165-167); body elongate and ... (Fig. 108, 109) .. [3] DYTISCIDAE narrowly subparallel (Fig.165-167); click mechanism —Antennae short, with a large pedicel; eyes each divided present, consisting of a prosternal process (Fig. 54, a) into two; forelegs long, raptorial; middle and hind legs which fits into a deep mesosternal cavity (Fig. 54, b); short and modified for swimming antennae filiform, serrate, pectinate, flabellate, or in- ... (Fig. 110) .. [4] GYRINIDAE crassate ... (p. 99) .. Group E
—97— —Pronotum and body shape not sas above; click mech- -Tarsal formula 2-2-2, 3-3-3, 3-4-4, 4-4-4, or 5-5-5, if anism absent; antennae often clubbed ... 11 4-4-4 then canthus absent and first 3 ventrites not fused together ... 17 11(10) Elytra short, truncate, exposing at least 2 ab- dominal tergites, usually more, the exposed tergites 17(16) Tarsal formula 5-5-5 (excluding those families well sclerotised rather than membranous; abdomen with 5-5-5-segmented tarsi already keyed out) usually flexible ... (p. 99) .. Group F ... (p. 102) .. Gr p —Elytra not so, broadly rounded posteriorly and usually —Tarsal formula 2-2-2, 3-3-3, 3 4 4, or 4-4-4; antennae covering entire abdomen, but if shorter then exposing filiform, moniliform, or with a distinct, compact, l-3- only the abdominal apex; tergites membranous or segmented club; mostly small beetles of average weakly sclerotised ... 12 length 2.5-3 mm, but some up to 17 mm long ... (p. 104) .. Gr p K 12(11) Antennae with an abrupt 3- or 4-segmented club, geniculate, usually retractable into a cavity on under- side of prothorax; body oval, disc-shaped, moderately Gr p A to strongly convex, glossy, usually glabrous and black; Includes families with a lamellate or flabellate (exception- elytra shortened, exposing the pygidium; length 2-9 ally pectinate) antennal club. (Fig. 114, 115) .. [6] IS E I AE —Antennae and body not as above ... 13 SCA A AEOI EA Lucanidae, Trogidae, Scarabaeidae. OS IC OI EA Bostrichidae (Euderia). Y OI EA 13(12) Body broadly oval, strongly convex, glossy; elytra Chelonariidae (Βrο n E E IO OI EA Rhipiphoridae. usually with 2 fine grooves near suture; in lateral view basal part of elytra and pronotum angular; antennae Al Lamellae of antennal club not capable of close with a 3-segmented club; tarsi 5-5-5- or 5-5-4-seg- apposition (Fig. 30) ... 2 mented but appearing 4-4-4-segmented because of —Lamellae of antennal club capable of close apposition vestigial segment 4 ... (Fig. 196) .. [48] A AC I AE (Fig. 31) .3 —Body not as above; elytra usually without 2 grooves near suture; in lateral view basal part of elytra and Α2( Antennae geniculate (exception: Ceratognathus pronotum not distinctly angular; antennae and tarsi not parrianus) ... (Fig. 139-143) .. [13] UCA I AE as above ... 14 —Antennae not geniculate ... 4
14(13) Frons with 1 or 2 ocelli ... (p. 100) .. Gr p G Α3(1 El tr d ll r r l t b r l t th t tr —Frons without ocelli ... 15 ... (Fig. 144) .. [14] OGI AE — El tr usually glossy, glabrous, or semiglabrous, or 15(14) Body loosely articulated, elongate, oblong to sub- with velvety pubescence (e.g., Costelytra), and often parallel; integument soft, often velvety in appearance with striae ... (Fig. 145-152) .. [15] SCA A AEI AE (e.g., Fig. 169, 186, 223), with decumbent pubescence and often additional scattered and erect setae, or with Α4(2 Metatarsi 4-segmented short and inconspicuous pubescence; pronotum and ... (Fig. 214) .. [64] I I O I AE elytra sometimes with ridges divided into cell-shaped - Metatarsi 5-segmented ... 5 compartments (Fig. 168) ... (p. 100) .. Gr p —Body compact; integument hard, glabrous or with Α5( Ant nn l l b th r th n 3 l ll ( p t 7 decumbent pubescence; pronotum and elytra not as ... (Fig. 164) .. [26] C E O A II AE (Brounia) above ... 16 —Antennal club of male with 3 flabella ... (Fig. 174) .. [35] OS IC I AE (Euderia) 16(15) Tarsal formula 5-5-4, exceptionally 4-4-4 in some Tenebrionidae, but then base of each antenna ob- scured from above by a shelf-like expansion (canthus Gr p Β [assisted by G. Kuschel] — Fig. 11, f) and first 3 ventrites fused together; body Includes families with tarsi 5-5-5-segmented but appear- broadly to narrowly oval, medium-sized to large (l.5- ing 4-4-4-segmented because of vestigial segment 4, and 20 mm); procoxae usually conical and projecting; segment 3 distinctly bilobed. Exception: Aglycyderinae antennae 11-segmented (10-segmented in Archaeo- (family Belidae) and Neocyba (Brentidae: Apioninae), in glenes, Tenebrionidae), filiform, serrate, incrassate, which tarsi appear 3-3-3-segmented but are 4-4-4-seg- sawtoothed, comblike, or flabellate, without a club or mented, and head with a rostrum. with a loosely defined, elongate, 3- or 4-segmented club ...(p. 101) .. Gr p I C YSOME OI EA Chrysomelidae ( n l Bruchinae), C r -
— 8— b d CU CU IO OI EA Nemonychidae, Anthribidae, ments combined, and ending in strong claws (exception: Belidae (incl. Aglycyderinae), Brentidae (incl. Apioninae), some Limnichidae ( ph l and some Byrrhidae, with Curculionidae (including Brachycerinae, Curculioninae, 4-4-4-segmented tarsi); body broadly oval and convex, Rhynchophorinae, Cossoninae, Scolytinae, Platypodinae). deflexed head concealed by pronotum from above, legs retractable into body cavities, prosternum with a process B1 Anterior part of head never with a rostrum 2 fitting into a cavity of mesosternum, and antennae with a —Anterior part of head elongate, forming a long or short weak club of 3-7 segments; aquatic or semi-aquatic. rostrum (exceptionally short in Scolytinae, Platy- podinae), with modified mouthparts (Fig. 6, 7) ... 3 Y OI EA Byrrhidae, Dryopidae, Elmidae, Limnichidae.
Β (1 Ant nn ll n rt d n fr nt l pr n n Cl Antennae shorter than pronotum, comblike, often nd p bl f b n fl x d b rd v r b d t concealed ... (Fig. 159) .. [21] YO I AE l t t -th rd l n b d —Antennae usually longer than pronotum, slender, saw- toothed, sometimes weakly clubbed ... 2 ... (Fig. 234-237) .. [76] CE AM YCI AE —Antennae not inserted on frontal prominences, not capable of being flexed backwards, usually only mod- C2(1) Body narrowly elongate; elytra with lateral mar- erately elongate (exception: Cryptocephalinae, with gins subparallel ... (Fig. 160) .. [22] E MI AE long antennae capable of being flexed backwards) —Body broadly oval; elytra with lateral margins broadly curved (Fig. 158, 161) .. 3 ... (Fig. 238-243) .. [77] C YSOME I AE
B3(l) Antennae geniculate, with a more or less compact C3(2) Hind margin of pronotum evenly curved or slightly club (Fig. 6, 28); scape long sinuate; frontoclypeal suture absent; body glabrous or semiglabrous ... (Fig. 158) .. [20] Y I AE ... (Fig. 251-260) .. [82] CU CU IO I AE —Antennae straight, with or without a club (Fig. 244- —Hind margin of pronotum strongly sinuate; fronto- clypeal suture present; body densely pubescent 249) ... 4 ... (Fig. 161) .. [23] IM IC I AE Β (3 br p r t d fr l p b d t n t t r (Fig. 6, b); mandibles with a mola 5 Gr p —Labrum fused with clypeus, without a suture; mandi- Includes families with hydrophilous species commonly bles without a mola bearing extremely elongate maxillary palps, usually longer than the antennae, and antennae 7-11-segmented with a Β5( r n t th t r n b l ; nt nn l - 2-5-segmented club. nt 3 r 5 xt nd n t ; p r pr nt n ll Y O I OI EA Hydrophilidae. S A Y I OI EA Hyd- tibiae; all 5 ventrites free raenidae. ... (Fig. 244) .. [78] EMO YC I AE —Pronotum usually with a transverse basal carina and a 1Body usually spherical or broadly oval and strongly lateral carina; antennal segment 1 or 2 extending to convex; maxillary palps usually shorter than antennae; eyes; spurs absent on all tibiae; only 5th ventrite antennae with a 3-segmented pubescent club completely free, other ventrites partly fused or firmly ... (Fig. 111-113) .. [5] Y O I I AE braced ... (Fig. 245, 246) .. [79] A I I AE —Body usually moderately narrowly elongate, rarely more broadly rounded; maxillary palps often longer Β ( tr d r t l l n t nd n rr ; pr t b than antennae (exception: M r p th species, with with a grooming device on inner face opposite tarsal maxillary palps much shorter than antennae); antennae articulation, consisting of fine and dense pubescence usually with a 5-segmented pubescent club in a broad, shallow groove or impression; gular sut- ... (Fig. 116, 117) .. [7] Y AE I AE ures distinctly separate or obsolete ... (Fig. 247, 248) .. [80] E I AE —Rostrum usually extremely elongate (Fig. 249); pro- Gr p Ε tibia without a grooming device; gular sutures merged Includes families with posterolateral angles of pronotum into one and always distinct strongly produced posteriorly, body narrowly elongate ... (Fig. 249, 250) .. [81] E I AE and subparallel, clicking apparatus present, and antennae filiform, serrate, pectinate, flabellate, or incrassate. Gr p C E A E OI EA Elateridae, Eucnemidae. Includes families with 5-5-5-segmented tarsi, each ter- minal segment enlarged, often longer than all other seg- Ε1 Labrum concealed beneath clypeus, externally not
-99 — visible; ventrites 4 and 5 connate 7( Antennae filiform, serrate, or pectinate but never ... (Fig. 165) .. [27] EUCNEMIDAE with a distinct club ... (Fig. 169) .. [30] CANTHARIDAE —Labrum visible; ventrites 4 and 5 not connate (in part; see also Group H) ... (Fig. 166, 167) .. [28] ELATERIDAE —Antennae with a distinct, compact club (Fig. 23) ... (Fig. 187) .. [42] NITIDULIDAE (in part)
Gr p Includes families with short and usually truncate elytra, Gr p G exposing at least 2 but usually more abdominal tergites, Includes families with frons bearing 1 or 2 ocelli. the exposed tergites well sclerotised, and abdomen usually flexible. Family Ptiliidae, with some species having trun- BOSTRICHOIDEA: Dermestidae (in part). DERODONTOIDEA: cate elytra and 2 abdominal tergites exposed, is included in Derodontidae. STAPHYLINOIDEA: Staphylinidae (Microsil- Group K (p. 104). phinae, Omaliinae, some Proteininae).
CHRYSOMELOIDEA: Chrysomelidae (Bruchinae). CUCUJOIDEA: G1 Head with 1 ocellus on frons, positioned medially Nitidulidae (in part; see also Group J), Monotomidae (in ... (Fig. 173) .. [34] DERMESTIDAE (in part) part; see also Groups J, K). ELATEROIDEA: Cantharidae (in ... [12] STAPHYLINIDAE (some Proteininae) part; see also Group STAPHYLINOIDEA: Staphylinidae —Head with 2 ocelli on frons, positioned laterally (Fig. (except Microsilphinae and some Proteininae of genus 123, 124) ... 2 Silphotelus). TENEBRIONOIDEA: Salpingidae (Inopeplinae only), Rhipiphoridae (see also Group A). G (l Head and pronotum not strongly sculptured; body smooth and glabrous or with sparse pubescence and 1 Elytra usually truncate posteriorly, exposing at least 3 punctation ... (Fig. 123, 124) .. [12] STAPHYLINIDAE but most often 6 or 7 abdominal tergites ... 2 (Microsilphinae, Omaliinae, some Proteininae) —Elytra truncate or rounded posteriorly, exposing 2 or —Head with a series of canals and bridges; pronotum rarely 3 abdominal tergites ... 4 divided by longitudinal ridges into several cell-like depressions ... (Fig. 170) .. [31] DERODONTIDAE F2(1) Abdomen short, broad, inflexible; head, pronotum, and often elytra with setose foveae .. (Fig. 126) .. [12] STAPHYLINIDAE (Pselaphinae) Gr p Η —Abdomen more elongate, narrow to moderately broad, Includes families with soft-bodied species having decum- flexible; usually 6 or 7 tergites exposed ... 3 bent pubescence and often additional scattered and erect setae, or with inconspicuous pubescence, and body sur- F3(2) Metatarsi 4-segmented; tarsal formula 5-5-4 face often with a velvety appearance. ... (Fig. 227) .. [72] SALPINGIDAE (Inopeplinae) — Metatarsi not 4-segmented, or if so then tarsal formula BYRRHOIDEA: Ptilodactylidae. CLEROIDEA: Cleridae, Mely- other than 5-5-4 ridae. ELATEROIDEA: Cantharidae (in part; see also Group ... (e.g., Fig. 125) .. [12] STAPHYLINIDAE (in part) F), Lycidae. EUCINETOIDEA: Scirtidae (=Helodidae). TENEBRIONOIDEA: Oedemeridae, Pyrochroidae, Scraptiidae. (1 Antennae appearing 10-segmented, with usually a 1-segmented club; elytra parallel-sided, usually with H1 Tarsal formula 5-5-5 ... 2 rows of punctures, truncate posteriorly, leaving pygid- —Tarsal formula 5-5-4 ... 7 ium exposed ... (Fig. 188, 189) .. [43] ΜΟΝΟΤΟΜΙ AΕ (Lenax, Monotoma) H2(1) Body flattened, loosely articulated; pronotum with —Antennae usually 11-segmented and not as above; several ridges dividing disc into irregularly shaped elytral shape not as above ... 5 cell-like depressions; elytra leathery, with strong, depressed punctures between longitudinal ridges 5( Tarsi appearing 4-4-4-segmented, with segment 3 ... (Fig. 168) .. [29] LYCIDAE bilobed; body robust, egg-shaped, usually broadest —Body form not as above; pronotum without ridges and towards apex; head often concealed from above irregularly shaped cell-like depressions; elytra hard, ... (Fig. 238) .. [77] CHRYSOMELIDAE (Bruchinae) with or without longitudinal ridges, but never with —Tarsi 5-5-5-segmented; body shape not as above ... 6 strongly impressed punctures between them ... 3
(5 Antennae flabellate; body wedge-shaped Η3( S t ll h rt- h p d; h nd r n f pr n t ... (Fig. 214) .. [64] RHIPIPHoRIDAE (see also Group A) th n t bl nt t th f r n tr n v r b —Antennae and body shape not as above ... 7 ( 1 3 [ 5] I O AC Y I AE
— 1 — —Scutellum subtriangular; hind margin of pronotum I1 Body moderately to strongly flattened (Fig. 192-195), without such teeth ... 4 often hairless; antennae filiform or beadlike, some- times with a loose 2-4-segmented club; mandibles Η (3 n lt t t r l nt pl n v r b l b d more or less pointing forwards; elytra usually striate ... (Fig. 186) .. [41] ME Y I AE (=Dasytidae) 2 ... (Fig. 183) .. [39] C E I AE (M t x n —Body and antennae not as above; mandibles usually —Penultimate tarsal segment bilobed ... 5 not pointing forwards; elytra striate or not
Η5( d ft n b l ndr l ll v r d th 12(1) Body moderately flattened, appearing pubescent; br tl h r ; h d br d br d pr n t r tarsal segment 2 and/or 3 lobed below or, if not lobed, br d r; pr n t h n rr r th n l tr ; nt- then antennal scape at least as long as 3 following seg- nn ft n th l 3- nt d l b ments combined; pronotum without sublateral ridges, ... (Fig. 184) .. [39] C E I AE often with anterior angles produced; antenna with or —Body flattened to moderately convex, lacking bristly without club (Fig. 192, 193) [45] SI A I AE hairs (pubescence short and decumbent); head usually —Body strongly flattened, usually glabrous (Fig. 194, narrower than pronotum; pronotum as broad as elytra 195); tarsal segments 2 and 3 not lobed below; anten- or slightly narrower; antennae without a 3-segmented nal scape as long as or shorter than the 2 following club ... 6 segments combined; pronotum usually with sublateral carinae; antennae without a club 3 Η (5 br l r t d nd d t n t v bl fr b v ; h d th h r t r t l b l ( 13( Pronotum without 2 sublateral carinae 1 ( 153 15 [1 ] SCI I AE ... (Fig. 194) .. [46] CUCU I AE —Labrum membranous and covered by clypeus, not —Pronotum with 2 sublateral carinae visible from above ... (Fig. 169) .. [30] CA A I AE ... (Fig. 195) .. [47] AEMO OEI AE
Η7(1 E n t pr j t n C- h p d tr n l r n t ; 1 (1 Base of each antenna covered by a canthus (Fig. 11, pronotum usually with 2 small basal impressions f); abdominal sternites 1-3 fused together ... (Fig. 233) .. [75] SC A II AE ... (Fig. 219-221) .. [68] E E IO I AE —Eyes prominent, slightly to strongly projecting, slight- —Base of antennae exposed; abdominal sternites 1-3 not ly emarginate or not; pronotum usually without 2 basal fused together ... 5 impressions ... 8 15(4) Pronotum distinctly narrower at base than elytra, Η (7 Eyes not strongly projecting laterally; only pen- half to two-thirds of elytral width (Fig. 228-232)... 6 ultimate tarsal segment bilobed; antennae usually fili- —Pronotum usually as wide at base as elytra, or nearly so form ... (Fig. 223) .. [70] OE EME I AE (exception: Ulodidae, with pronotum much narrower —Eyes strongly projecting laterally; penultimate and basally than elytra– Fig. 217) ... 9 antepenultimate tarsal segments lobed; antennae usu- ally pectinate or serrate 1 (5 Pronotum without lateral margins; body ant-shaped ... (Fig. 224, 225) .. [71] Y OC OI AE (Fig. 228-232) ... 7 —Pronotum with lateral margins; body not as above ... 8
Gr p I I7(6) Ventrites all free; penultimate tarsal segment lobed; eyes with fine facets, not notched near antennal base; Includes families with 5-5-4-segmented tarsi, exceptionally 4-4-4-segmented in some Tenebrionidae (but then base of elytral epipleura incomplete antenna covered by a canthus), procoxae usually conical ... (Fig. 228-231) .. [73] A ICI AE —Basal 2 ventrites connate; penultimate tarsal segment and projecting, and antennae without or with a loosely defined, elongate, 3- or 4-segmented club. small, simple, and antepenultimate segment lobed beneath; eyes coarsely faceted, slightly notched near CUCU OI EA Cucujidae, Laemophloeidae, Silvanidae (in antennal base; elytral epipleura absent part; see also Group E E IO OI EA Aderidae, ... (Fig. 232) .. [74] A E I AE (= Euglenidae) Anthicidae (incl. Pedilinae), Archeocrypticidae, Chalco- dryidae, Melandryidae, Mordellidae, Phloeostichidae 1 ( Pronotum and elytra with fine and dense punc- (males only; females in Group F), Salpingidae (excl. tation; antennae slightly incrassate, but without a Inopeplinae: see Group F), Tenebrionidae, Ulodidae distinct club; body form not as below (formerly as Zopheridae). ... (Fig. 190) .. [44] OEOS IC I AE (males of A p th see also Group J)
-1 1- —Pronotum usually coarsely punctate, and elytra with Chaetosomatidae, Phycosecidae, Trogossitidae (incl. rows of coarse punctures; antennae with a 3-seg- Protopeltinae). CUCUJOIDEA: Cavognathidae, Cryptophag- mented, loosely formed club; often resembling small idae (in part; males with tarsi 5-5-4 are in Group I), carabid beetles ... (Fig. 226) .. [72] SALPINGIDAE Cucujidae, Laemophloeidae, Silvanidae (in part; see also (excl. Inopeplinae) Group I), Erotylidae, Languridae, Nitidulidae (in part; see also Group F), Phloeostichidae, Monotomidae (in part; I9(5) Antennae with a 3-segmented, elongate club ... 10 see also Groups F and K). EUCINETOIDEA: Eucinetidae. —Antennae filiform, serrate, or slightly incrassate, with- STAPHYLINOIDEA: Agyrtidae, Leiodidae, Scydmaenidae. out a distinct 3-segmented club ... 11 J1 Pronotum hood-like in lateral view, partially covering I10(9) Body irregularly oval in outline; pronotum and head from above; antennae, if clubbed, then with all elytra with margins not as below segments long and club not symmetrical; body shape ... (Fig. 191) .. [44] PHLOEOSTICHIDAE from elongate cylindrical to oval (males of Priasilpha) ... (Fig. 177) .. [36] ANOBIIDAE (excluding Ptininae) —Body approximately oval in outline; pronotum and —Pronotum not as above; antennae, if clubbed, then with elytra with distinct sharp margins club symmetrical (exception: some Dermestidae); ... (Fig. 210) .. [60] ARCHEOCRYPTICIDAE body shape rarely cylindrical
I11(9) Body broadly oval, flattened, with decumbent or J2(1) Antennae not clubbed 3 erect pubescence, bristles, or scales forming patterns; —Antennae with a more or less distinct, 1-8-segmented, pronotum narrower than elytral base, often with api- elongate and loosely formed or compact club; if club cal angles produced ... (Fig. 217) .. [66] ULODIDAE not distinct then segment 8 smaller than adjoining (formerly as Zopheridae) segments —Body narrowly elongate, convex to slightly flattened, with decumbent pubescence often forming patterns; J3(2) Body appearing spider-like; legs and antennae long pronotum as wide as elytral base, with apical angles and thin; antennal insertions close together; body rounded but not produced ... 12 covered with dense pubescence or scales forming patterns ... (Fig. 178) .. [36] ΑΝΟΒΙΙ ΑΕ I12(11) Body narrowly subparallel, flattened; head not (Ptininae) deflexed, not deeply inserted into prothorax; tibiae —Body elongate or oval; legs and antennae moderately without combs or spurs long; antennal insertions distant from each other ... 4 ... (Fig. 218) .. [67] CHALCODRYIDAE —Body wedge-shaped, tapering posteriorly, convex; J4(3) Body elongate-oval, narrowed posteriorly; head head deflexed, deeply inserted into prothorax; tibiae concealed from above, resting against procoxae in nat- often with serrate combs or spines (Fig. 84) ... 13 ural position; metacoxae greatly expanded, covering much of 1st ventrite; elytra often with cross-striations I13(12) Last sclerotised tergite usually covered by elytra ... (Fig. 155) .. [ 17] EUCINETIDAE and not forming an acute process extending beyond —Body broadly to narrowly elongate, subparallel, often elytra; head not abruptly constricted behind eyes; flattened; head usually prognathous, exposed and not pronotum with lateral carinae incomplete; metacoxae concealed by pronotum; metacoxae not enlarged; not plate-shaped ... (Fig. 212) .. [62] MELANDRYIDAE elytra without cross-striation ... 5 —Last sclerotised tergite produced posteriorly and forming an acute process extending beyond elytra; J5(4) Body with long and erect hairs, sparsely and head abruptly constricted behind eyes; pronotum with coarsely punctate, moderately flattened; elytra not lateral carinae complete; metacoxae enlarged, plate- striate ... (Fig. 182) .. [38] CHAETOSOMATIDAE shaped ... (Fig. 213) .. [63] MORDELLIDAE —Body without long and erect hairs, glabrous or with short, sparse pubescence and coarse punctation; elytra usually striate ... 6 Group J Includes families with tarsal formula clearly 5-5-5 not J6(7) Body moderately flattened, usually pubescent; already listed (see Groups B, C, D, F, and H). tarsal segment 2 and/or 3 lobed below or, if not lobed, then antennal scape at least as long as the 3 following BOSTRICHOIDEA: Anobiidae, Bostrichidae (in part, incl. segments combined; pronotum without sublateral car- Lyctinae; see also Group A), Dermestidae (in part; see also inae ... (Fig. 192, 193) .. [45] SILVANIDAE Group G), Jacobsoniidae (in part; see Group K for species —Body strongly flattened, appearing glabrous; tarsal with tarsi 3-3-3), Nosodendridae, Ptinidae. CLEROIDEA: segments 2 and 3 not lobed below; antennal scape as
-102- long as or shorter than the 2 following segments J14(13) Body surface glabrous or with inconspicuous combined ... 7 pubescence, usually strongly glossy ... 15 —Body surface moderately to densely pubescent, matt or J7(6) Pronotum without 2 sublateral carinae slightly glossy ... 18 ... (Fig. 194) .. [46] CUCUJIDAE —Pronotum with 2 sublateral carinae J15(14) Body narrowly elongate, subparallel to cylindri- ... (Fig. 195) ... [47] AEMO OEI AE cal; pronotum slightly to distinctly elongate ... 16 —Body broadly to narrowly oval; pronotum transverse J8(2) Antennae with a 3-5- or 7-segmented club and and narrow, as wide as elytral base ... 17 usually with segment 8 smaller than adjoining ones (Fig. 24), or if not so then body densely pubescent; J16(15) Body narrowly elongate, not cylindrical, sub- often glabrous beetles; legs usually bearing strong parallel; pronotum without rasplike teeth at front; spines ... (Fig. 120, 121) .. [10] EIO I AE metasternum extremely elongate, as long as all 5 ven- —Antennae usually with segment 8 normally developed; trites combined or longer; body length 0.8-2 mm usually pubescent beetles; legs without spines or with ... (Fig. 171) .. [32] ACO SO II AE small spines ... 9 —Body cylindrical or narrowly elongate and slightly flattened; pronotum often with rasplike teeth at front; J9(8) Body oblong to broadly oval, robust, with short, antennal club often asymmetrical (enlarged to one dense pubescence or with erect hairs or scales often side); metasternum not extremely elongate, much forming patterns; lower surface often with close, silky shorter than all 5 ventrites combined; body length 2.8- pubescence; elytra if striate then with a scutellary 6 mm .... (Fig. 174-176) .. [35] OS IC I AE striole; antennae with club 3-8-segmented, usually 3- (incl. Lyctinae; see also Group A) segmented ... (Fig. 173) .. [34] E MES I AE —Body shape variable, glabrous or with sparse to dense J17(15) Body black, broadly oval, strongly convex; pubescence, rarely forming patterns (with patterns in elytra with tufts of short pubescence in rows; legs Trogossitidae, some Nitidulidae); lower surface with- retractable into ventral body cavities; scutellum large, out silky pubescence; elytra with or without striae; triangular ... (Fig. 172) .. [33] OSO E I AE scutellary striole absent ... 10 —Body often bicoloured, narrowly oval, moderately convex to slightly flattened, glabrous; legs not J10(9) Antennal club 5-segmented; body flat, strongly retractable into ventral body cavities; scutellum small glossy; elytra with longitudinal striae ... (Fig. 200) .. [52] E O Y I AE ... (Fig. 119) .. [9] AGY I AE —Antennal club 1-3-segmented ... 11 J18(14) Elytral shoulders more or less produced ant- eriorly; elytral suture with a posterior gap (Fig. 93) J11(10) Antennal club 1- or 2-segmented; anterior pro- created by widened sutural flanges; pronotum with notal angles produced (Fig. 185, 189) ... 12 indistinct and incomplete lateral carinae; frons usually —Antennal club 3-segmented; anterior pronotal angles with a pair of cavities; basal antennal segment enlar- not produced ... 13 ged, as long as the 2 following segments combined ... (Fig. 197) .. [49] CAVOGNATHIDAE J12(11) Body elongate-oval; pronotum horseshoe shaped, —Elytral shoulders not produced anteriorly; elytral with anterolateral angles strongly produced anteriorly; suture complete (Fig. 94) (exception: Cryptophagidae); pygidium covered by elytra; tarsi pubescent pronotum usually with complete lateral carinae; frons ... (Fig. 185) .. [40] YCOSECI AE without cavities; basal antennal segment slightly en- —Body narrowly elongate; pronotum oval or rectangu- larged, usually shorter than segments 2+3 ... 19 lar, with anterolateral angles more or less produced; pygidium exposed; tarsi glabrous; head with promi- J19(18) Body narrowly subparallel, subcylindrical to nent temples ... (Fig. 188, 189) .. [43] ΜΟΝΟΤΟΜΙ ΑΕ slightly flattened, moderately elongate; eyes coarsely ( n x M n t faceted; head, pronotum, and elytra nearly equal in width; elytra striate or seriate-punctate; tarsal seg- J13(11) Body ant-like; antennal club loose; dorsal sur- ments 2 and 3 with setose lobes face usually with long pubescence; head with a distinct ... (Fig. 199) .. [51] A GU II AE neck, nearly as wide as pronotum; pronotum and elytra — d broadly to narrowly oval, rarely subparallel; eyes oval, with humeri rounded finely to moderately coarsely faceted; head distinctly ... (Fig. 122) .. [11] SCY MAE I AE narrower than pronotum; elytra as broad as pronotum, —Body not ant-like; antennal club distinct, usually com- usually without distinct striae; tarsal segments 2 and 3 pact; dorsal surface glabrous or variably pubescent...14 without setose lobes ... 20
-1 3--1 3- J20(19) Antennae usually closely approximated at base, 3-3-3; antennae 10- or 11-segmented, incrassate or with a often diverging to form a distinct V; pronotum often l-3-segmented club), Mycetophagidae (tarsi 3-4-4 or 4-4- depressed at base; body often with silky pubescence 4; antennae 11-segmented with a 3- or 4-segmented club), ... (Fig. 198) .. [50] C Y O AGI AE Prostomidae (tarsi 4 4 4; antennae 11-segmented with a 3- —Antennal bases broadly separated, and antennae not segmented club). forming a V-shape; pronotum not depressed; body with short pubescence and sometimes with additional Κ1 r l f r l - - ; b d n t ft n l th n 1 longer hairs ... 21 l n br dl t n rr l v l; l tr ft n x- p n 1 r bd n l t r t ; h nd n f th r- J21(20) Antennal club abruptly compact or ball shaped; l fr n d th l n h r ; nt nn th n b r n tarsal segments 1-3 more or less dilated, and segment l n h r nd th - r 3- nt d l b 4 sometimes smaller ... (Fig. 118) .. [8] I I AE ... (Fig. 187).. [42] I I U I AE (in part) —Tarsal formula 3-3-3, 3-4-4, or 4-4-4; body usually —Antennal club loosely formed, not ball shaped; tarsal over 1 mm long, variable in shape; elytra covering segments simple ... 22 abdomen (exception: Rhizophaginae); hind wings not featherlike, not fringed with hairs ... 2 J22(21) Body regularly oval and convex or elongate and Κ (1 d br dl v l t h ph r l; h d r r subparallel; pronotal base usually not sinuate; pub- l n l d b pr n t ... 3 escence short or long, with bristles or scales often —Body oblong oval to narrowly elongate and sub- forming patterns parallel; head not concealed by pronotum ... 5 ... (Fig. 179-181) .. [37] OGOSSI I AE —Body irregularly oval and flattened, narrowed post- Κ3( r - - nt d th ll nt pl ; eriorly; pronotal base sinuate; pubescence sparse and nt nn -1 - nt d th - nt d l b; short, grouped into clumps on elytra t x xp nd d nt l r pl t n l n f - OEOS IC I AE (Priasilpha) ... (Fig. 191) .. [44] r ; p rtl r nt r l d v d d; b d ll ( 9- 1 5 n l f p bl f r ll n nt b ll ... (Fig. 156) .. [18] C AM I AE Gr p Κ —Tarsi truly 3-3-3-segmented or appearing 3-3-3-seg- Includes families with tarsal formula 2-2-2, 3-3-3, 3-4-4, mented, with at least the 2nd bilobed; antennae 7-11- or 4-4-4; antennae often with a 1-3-segmented distinct segmented with a 1-6- (most often 3)-segmented club; club; mostly small beetles averaging 2.5-3.0 mm, though metacoxae without large plates; eyes not divided ... 4 some species up to 17 mm long. Κ (3 d p rt ll r nt r l n l d b pr n t OS IC OI EA Jacobsoniidae (see also Group J) (tarsi 2- b t nt r r p rt f pr n t ll r n t nd 2-2, or 5-5-5 but appearing 3-3-3; antennae 10-or 11-seg- n t xt nd d nt h rp r n v r h d; b d mented with a 1-3-segmented club). Y OI EA Hetero- ft n l r r (1-7 ft n p tt d ll l br ceridae (tarsi 4 4 4; antennae 11-segmented with a 6- or 7- r b l br ; nt nn h rt r th n pr n t th segmented club). CUCU OI EA Bothrideridae (tarsi 4-4-4; 1- - nt d l b; x ll r p lp r f r r antennae 11-segmented with a 2- or 3-segmented club), th p x bl l tr n t ( 13-15 Cerylonidae (tarsi 3-3-3 or 4 4 4; antennae 11-segmented ... (Fig. 206) .. [56] COCCI E I AE with a l- or 2-segmented club), Coccinellidae (tarsi 3-3-3 —Head partially or entirely concealed by pronotum, with or 4-4-4; antennae 7-11-segmented, incrassate or with a anterior part of pronotum extending into a sharp 1-6-segmented club), Corylophidae (tarsi 3-3-3; antennae margin over head; body small (0.5-2.5 mm), usually 11-segmented with a 3-segmented club), Endomychidae unicoloured, densely pubescent; antennae longer than (incl. Holoparamecinae) (tarsi 3-3-3; antennae 11-seg- pronotum, with a 3-segmented club; maxillary palps mented with a 2- or 3-segmented club), Corticariidae not as above ... (Fig. 207) .. [57] CO Y O I AE (=Lathridiidae) (tarsi 3-3-3; antennae 11-segmented with a 2- or 3-segmented club), Monotomidae (see also Groups Κ5( d xp d; nt nn 1 - nt d th F and J) (tarsi 4 4 4; antennae 10-segmented with a 1- or 2- l- r - nt d l b; t r l f r l - - segmented club). EUCI E OI EA Clambidae (tarsi 4 4 4; ... (Fig. 188, 189) .. [43] MO O OMI AE antennae 8-10-segmented with a 2-segmented club). (Lenax, Monotoma) S A Y I OI EA Ptiliidae (tarsi 2-2-2; antennae 10- or 11- —Pygidium covered by elytra or (Prostomidae) partially segmented with a2- or 3-segmented club). E E IO OI EA exposed; antennae usually 1 1-segmented (10-segmen- Ciidae (tarsi 4 4 4 or 3-3-3; antennae 8-10-segmented ted in some Colydiidae), with a 1-4-segmented club; with a 2- or 3-segmented club), Colydiidae (tarsi 4-4-4 or tarsal formula 3-3-3, 3-4-4, or 4-4-4 ... 6
-104- Κ (5 Ant nn l n dth f l p th —Body elongate to broadly oval, flattened or moderately pectinate 7-segmented club; tibiae flattened, spin-ose; convex, rarely cylindrical (exception: some body robust; elytra subparallel, with paler longi- Colydiidae), variably pubescent, rarely with bristles; tudinal spots ... (Fig. 162) .. [24] ΗΕΤE OCΕ 1 ΑΕ head and pronotum without horns; head usually not —Antennae much longer than width of clypeus, with a 1- deflexed ... 14 4-segmented club; tibiae not spinose; body usually not robust; elytra rarely with spots ... 7 Κ1 (13 Ant nn l n rt n xp d; nt nn l l b f- r - nt d; b d ll n rr l l n t nd Κ7( Ant nn th l l b; t pl tr n l pr - l htl fl tt n d th d r l rf l br nd d d p t r rl ; pr n nt pr j t n ; b d l r th p r h rt br tl ; l tr tr t tr n l d pr d bp r ll l - l n ... (Fig. 203) .. [54] CE Y O I AE ... (Fig. 222) .. [69] OS OMI AE —Antennal insertions concealed from above; body with —Antennae with a strongly defined, usually compact dorsal surface usually pubescent ... 15 club; temples not produced posteriorly; eyes moder- ately to strongly prominent; body convex to slightly Κ15(1 r l f r l 3- - ( l r - - ; d f depressed, subparallel to broadly oval ... 8 pr n t thl nt n th d f l tr ; pr n t ft n th d pr n n r b ; b d Κ (7 r n t d t n tl n rr r th n l tr l b br dl t n rr l v l d n l p b nt ft n (exception: th t n , Corticariidae) ... 9 th r n r ll h p t n l tr r n f r l —Pronotum as broad as elytral base or slightly narrower br n; nt nn ll th l 3- r - - ... 12 nt d l b; nt nn l n rt n xp d ... (Fig. 209) .. [59] MYCE O AGI AE Κ9( r l f r l - - ; nt nn l n rt n xp d; —Tarsal formula 3-3-3 or 4-4-4 in both sexes; sides of nt nn th p t - nt d l b; pr n t pronotum not as above; pronotum without 2 basal ll th d l pr n; l tr th d t n t depressions and differently sculptured, often covered tr ; b d n rr l v l d pr d l with pubescence or scales forming patterns; antennae ... (Fig. 201, 202) .. [53] O I E I AE with an abrupt, 1-3-segmented club; antennal inser- —Tarsal formula 3-3-3 ... 10 tions concealed ... (Fig. 215, 216) .. [65] CO Y II AE
Κ1 (9 r n t d t n tl t d th b l r f l t r ll l n t f v ; nt nn th - - nt d ph r l l b nd th nt pp r n t b f d t th r; ll b tl 1 -1 7 l n ... (Fig. 205) .. [55] E OMYC I AE (Holoparamecinae) —Pronotum not as above ... 11
Κ11(1 d l th n l n r t-br n; h d br ptl n tr t d b h nd ; nt nn th l b pp r n 1- nt d [3 ] ACO SO II AE ( r l th (see also Group J) Body 0.8-3.0 mm long, brown to black; head not abruptly constricted behind eyes; antennae with club 2- or 3-segmented ... (Fig. 208) .. [58] CO ICA II AE (= Lathridiidae)
Κ1 ( r n t th n p r d h rp r n n th r d f d ( x pt n l p r n ( [55] E OMYC I AE —Pronotum not as above ... 13
Κ13(1 d l n t ll l ndr l l br r th r t br tl ; pr n t nd/ r h d ft n b r- n t b r l r h rn ; h d d fl x d; nt nn l l b l r l - r 3- nt d; b d l n th 1-3 ( 11 [ 1] CII AE
— 1 5 — II: I EA KEY 8(5) Tarsi pseudotetramerous (5-5-5 but appearing 4-4- [pr v d d b .C. W tt] 4), with segment 3 usually strongly bilobed, rarely Families with tarsal formula 5-5-5 in females but 5-5-4 in some or pseudotrimerous (4-4-4 but appearing 3-3-3) (Fig. 67-81) ... CHRYSOMELOIDEA, CURCULIONOIDEA, all males—MONOTOMIDAE, CUCUJIDAE, LAEMOPHLOEIDAE, BYRRHOIDEA.. CRYPTOPHAGIDAE (all N.Z. genera), PHLOEOSTICHIDAE (all N.Z. some 9 genera), and PHALACRIDAE (Cyclaxyra) —key out in two places. —Tarsi not so ... 19
1 Metacoxae large, fused to metasternum, completely div- 9(8) Antennae lamellate (Fig. 22); pronotum humped iding 1st ventrite (Fig. 88, 89); sternites 1-3 fused; laterally ... (Fig. 164) .. [26] CHELONARIIDAE prothorax with distinct notopleural sutures (Fig. 2) —Antennae and pronotum not so ... 10 ... ADEPHAGA .. 2 —Metacoxae varying in size, usually movably articu- 10(9) Head without a rostrum, or rarely (Acanthoscelides) lated to metasternum, never completely dividing 1st slightly rostrate; antennae without a club, not geni- ventrite (Fig. 90); all sternites usually free; prothorax culate; antennal scrobes absent; gular sutures distinct without notopleural sutures (Fig. 3) .. POLYPHAGA .. 5 and separate ... CHRYSOMELOIDEA .. 11 —Head usually produced into a rostrum (Fig. 6, 7) 2(1) Hind legs without swimming hairs; prosternum not (rostrum short and not apparent in Scolytinae, Platy- prolonged behind as a median keel; terrestrial beetles podinae); antennae with a more or less distinct club, not streamlined for aquatic life ... 3 often geniculate and retractable into scrobes on side of —Hind legs with swimming hairs; prosternum prolonged rostrum (Fig. 6, d); gular sutures usually confluent behind as a median keel; body streamlined for aquatic (Fig. 7) or obsolete ... CURCULIONOIDEA.. 13 life ... 4 11(10) Antennae inserted on tubercles, and capable of 3(2) Antennae moniliform (Fig. 19); head and pronotum being flexed backwards against body, usually extend- with deep, paired, longitudinal grooves ing at least to base of elytra ... (Fig. 101) .. [1] RHYSODIDAE ... (Fig. 234-237) .. [76] CERAMBYCIDAE —Antennae usually filiform (Fig. 1), never moniliform; —Antennae not inserted on tubercles, not capable of head and pronotum without deep, paired, longitudinal being flexed backwards against body, usually not grooves ... (Fig. 102-107) .. [2] CARABIDAE extending to base of elytra ... 12 (incl. Cicindelinae) 12(11) Head somewhat rostrate; antennae and body 4(2) Eyes not divided, or absent (e.g., Kuschelydrus); bearing scales ... (Fig. 238) .. [77] CHRYSOMELIDAE hind legs longer than forelegs or middle legs (Bruchinae) ... (Fig. 108, 109) .. [3] DYTISCIDAE —Head not at all rostrate; antennae and body without —Eyes completely divided into upper and lower parts; scales ... (Fig. 239-243) .. [77] CHRYSOMELIDAE forelegs distinctly longer than middle or hind legs (excl. Bruchinae) ... (Fig. 110) .. [4] GYRINIDAE 13(10) Antennae usually geniculate (Fig. 6, 28), with 1st 5(1) Antennae with 10 or fewer segments, the terminal segment retractable into scrobes (Fig. 7, d) segments lamellate, produced anteriorly into laterally ... (Fig. 251-260) .. [82] CURCULIONIDAE flattened plates (Fig. 30, 31) ... SCARABAEOIDEA .. 6 —Antennae not geniculate ... 14 —Antennae rarely lamellate or flabellate, if so then 11- segmented ... 8 14(13) Tarsi appearing 3-3-3; body form characteristic, as figured .. (Fig. 248) .. [80] BELIDAE (Aglycyderinae) 6(5) Antennae geniculate, with club segments not able to —Tarsi appearing 4 4body-4; form not so ... 15 be folded closely together; mandibles prominent ... (Fig. 139-143) .. [13] LUCANIDAE 15(14) Labrum distinct, separate (Fig. 6, b) ... 16 —Antennae not geniculate, with club segments able to be —Labrum not distinct, not separate ... 17 folded closely together; mandibles not prominent ... 7 16(15) Prothorax without transverse or lateral carinae; 7(6) Abdomen with 5 ventrites; elytral sculpture very abdominal ventrites free and not strongly sclerotised rough ... (Fig. 144) .. [14] TROGIDAE ... (Fig. 244) .. [78] NEMONYCHIDAE —Abdomen with 6 ventrites; elytra smooth, or if —Prothorax with a transverse carina and often additional sculptured then not rough lateral carinae; ventrites 1-4 fused, strongly sclero- ... (Fig. 145-152) .. [15] SCARABAEIDAE tised ... (Fig. 245, 246) .. [79] ANTHRIBIDAE
—106—
17(15) Body strongly elongate, at least 18 mm long in- abdominal ventrites free; body form not as above; hind
cluding rostrum (Fig. 249) [81] BRENTIDAE angles of pronotum at most rectangular, not produced (Brentinae) backwards 24
—Body length considerably less than 18 mm 18 23(22) Labrum not visible externally; antennal insertions
18(17) Trochanters strongly oblique, not separating fem- distant from eyes (Fig. 165) .. [27] EUCNEMIDAE ora and coxae (Fig. 62); 2 gular sutures, distinct and —Labrum free, visible externally; antennae inserted near
separate (Fig. 247) .. [80] BELIDAE (Belinae) eyes (Fig. 166, 167) .. [28] ELATERIDAE —Trochanters slightly oblique, distinctly separating femora and coxae (Fig. 63); 1 gular suture, positioned 24(22) Minute beetles less than 1 mm long; antennae medially (Fig. 250) .. [81] BRENTIDAE (Apioninae) slender, filiform, with last 3 segments enlarged; wings reduced to a central axis with anterior and posterior
fringes of long hairs (Fig. 118) .. [8] PTILIIDAE 19(8) Antennae geniculate, with a compact 3-segmented —Larger beetles, length exceeding 1 mm; antennae not club; elytra truncate, exposing last 2 broad abdominal so; wings (if present) with a broad lamina and venation
tergites, never regularly 9-or 10-striate; body strongly 25 sclerotised, usually black, distinctive in form (Fig. 114, 115) .. [6] HISTERIDAE 25(24) Elytra truncate, leaving usually 6 sternites ex-
—Antennae rarely geniculate, or if so then without a posed (Fig. 124, 127-130) .. [12] STAPHYLINIDAE compact and 3-segmented club; other characters never (in part)
all present in combination 20 —Elytra rarely truncate, leaving fewer than 6 sternites
exposed (usually 1-3) 26 20(19) Metacoxae with posterior face vertical and at least slightly, usually strongly, excavate to receive retracted 26(25) Loosely articulated, elongate beetles with soft femur (Fig. 66); antennae filiform, serrate, pectinate, integument (e.g, Fig. 169); elytra not closely em- or thickened but never with a true club; ocelli absent; bracing abdomen laterally; metacoxae very short,
procoxal cavities open behind (Fig. 40) 21 without plate-like extensions of ventral hind margin
—If metacoxae with posterior face vertical and excavate (femoral plates) 27 (Nosodendridae, Dermestidae, Anobiidae) then anten- —Compact, usually less elongate beetles, with firmer nae with a distinct 3-segmented club, or head with a integument; elytra closely embracing abdomen later- median ocellus, or prothorax produced over head in ally; metacoxae usually longer, and with distinct
the form of a hood; procoxal cavities usually partly or femoral plates (Fig. 52, 53) 28
completely closed behind (Fig. 41) 37 27(26) Elytra with longitudinal costae between pairs of 21(20) Abdomen with 2 basal ventrites fused together, striae; trochanters long, transversely joined to femora
the suture partly obliterated; tarsi with ventral adhes- (Fig. 64) (Fig. 168) .. [29] LYCIDAE ive lobes on segments 2-4; body form distinct, cylin- —Elytra without distinct striae or longitudinal costae;
drical anteriorly (Fig. 157) .. [33] BUPRESTIDAE trochanters short, obliquely joined to femora (Fig. 65)
—Abdomen with all ventrites usually free, or if fused (Fig. 169) .. [30] CANTHARIDAE (Elateroidea) then suture between 1st and 2nd vent- rites as distinct as that between 2nd and 3rd; tarsi 28(26) Base of pronotum crenulate; scutellum heart
rarely with adhesive lobes on more than 1 segment; shaped (Fig. 163) .. [25] PTILODACTYLIDAE
body form not as above 22 —Base of pronotum not crenulate; scutellum not so 29
22(21) Anterior median part of mesosternum deeply and 29(28) Metacoxae at least narrowly separated by a tri- narrowly excavate, with sides of cavity vertical, angular anterior median projection of basal sternite; receiving narrow, pointed posterior process of body without sessile scales; anterior part of proste- prosternum, these together usually forming part of rnum somewhat produced forwards to partly cover
`click mechanism' (Fig. 54); abdomen with basal 4 ventral Surface of head Y OI EA (in part) .. 30 ventrites fused; body form characteristic, as in Fig. —Metacoxae contiguous, or if narrowly separated 165-167; hind angles of pronotum almost always (Byrrhidae, Curimus) then body clothed dorsally with
produced backwards, partly around elytral shoulders sessile scales; prosternum emarginate anteriorly 34
ELATEROIDEA.. 23 —Anterior median part of mesosternum shallowly and 30(29) Antennae very short, with anterior processes on broadly excavate, or not excavate at all; posterior last 7 or 8 segments (Fig. 29); body densely setose or
prosternal process absent, or not shaped as above; with very long pubescence (Fig. 159, 162) 31
—107— —Antennae not as above; body neither densely setose —Antennae not as above, longer than maxillary palps; nor with very long pubescence ... 32 head without a Y-shaped impressed line ... 39
31(30) Legs without spines; tarsal formula 5-5-5 38(37) Body usually very convex, elongate-oval to ... (Fig. 159) .. [21] YO I AE almost hemispherical; abdomen with 5 ventrites —Legs, especially forelegs, with a row of spines on ... (Fig. 111-113) .. [5] Y O I I AE external edge; tarsal formula 4-4-4 —Body less convex, more elongate; abdomen with 6 or 7 ... (Fig. 162) .. [24] ΗΕ Ε OCΕ Ι ΑΕ ventrites ... (Fig. 116, 117) .. [7] Y AE I AE 32(30) Metacoxae widely separated, oval; body less than 39(37) Elytra truncate at apex, leaving at least 3 sclero- 1.4 mm long; intertidal ... [23] IM IC I AE (Hyphalinae) tised abdominal tergites uncovered; antennae filiform —Metacoxae narrowly separated, transverse; body or thickened towards apex, but without a strong, larger, length more than 1.4 mm; not intertidal ... 33 compact club (exception: Staphylinidae : Euaesthet- inae, Microsilphinae) ... 40 33(32) Body elongate, with pubescence not forming a —If 3 or more sclerotised tergites exposed then elytra not pattern; ventral surface without grooves for legs and truncate (Salpingidae : Inopeplinae), or antennae with a strong, compact club (Nitidulidae) ... 41 antennae ... (Fig. 160) .. [22] E MI AE —Body broadly oval, with pubescence forming a pattern; legs and antennae retractable into grooves on ventral 40(39) Abdomen with very limited dorsoventral flexibility; surface... (Fig. 161).. [23] IM IC I AE (Limnichinae) maxillary palps usually long and modified (Fig. 16); integument with characteristic deep foveae in various 34(29) Metacoxae scarcely contiguous, occasionally not positions, especially on vertex of head and pronotum contiguous; body broad, convex; mesocoxae widely ... (Fig. 126) .. [12] S A Y I I AE (Pselaphinae) separated (Fig. 52, 53); legs and antennae retractable —Abdomen flexible dorsoventrally; maxillary palps into grooves or depressions on ventral surface usually moderately long and not modified; integument ... (FIg. 158) .. [20] Y I AE rarely with such foveae —Metacoxae strongly contiguous; body form variable, ... (e.g., Fig. 127-138) .. [12] S A Y I I AE not as above; mesocoxae narrowly separated; legs and (excl. Pselaphinae, Silphotelus) antennae usually not retractable into grooves or de- pressions on ventral surface ... EUCIΝΕ OI EA 35 41(39) Body less than 1.8 mm long, convex; vertex with a pair of deep foveae between eyes; sides of frons with 35(34) Metacoxae considerably narrowed laterally; fem- a deep semicircular incision in front of each eye oral plates small; 4th tarsomere lobed below; head ... (Fig. 125) .. [12] S A Y I I AE (Silphotelus) with characteristic keels behind and below eyes (Fig. —Head without this combination of foveae and lateral 12) ... (Fig. 153, 154) .. [ 16] SCI I AE incisions ... 42 —Metacoxae not narrowed laterally; femoral plates very large; tarsi without lobed segments; head without such 42(41) Metacoxae with posterior face vertical and ex- keels ... 36 cavated to receive retracted femur (Fig. 56) ... 43 —Metacoxae with posterior face not vertical ... 47 36(35) Elongate-oval beetles; head narrow, concealed by pronotum; tarsal formula 5-5-5 43(42) Head with a pair of ocelli between and close to ... (Fig. 155) .. [17] EUCI E I AE eyes; small beetles, length less than 1.8 mm —Hemispherical beetles; head very broad, reflexible ... (Fig. 170) .. [31] E O O I AE against a cavity (bounded posteriorly by a semicircular —Paired ocelli absent ... 44 carina) on underside of thorax; tarsal formula 4-4-4 ... (Fig. 156) .. [18] C ΑΜΒΙ AE 44(43) Head with a median ocellus (sometimes obscured by long, coarse setae) ... [34] E MES I AE (in part) —Head without any ocellus ... 45 37(20) Antennae short, with 6th segment modified as a cupule (Fig. 26, 27) and terminal 3 or 4 segments 45(44) Prothorax produced over head in a hood-like forming a strong, pubescent club (exception: Meropa- fashion; antennae serrate, flabellate, or with 3 seg- thus, with club elongate and not as above — Fig. 117), ments each enlarged but not forming a compact club or, if club weak and not pubescent, then maxillary ... (Fig. 177) .. [36] A O II AE (Anobiinae) palps much longer than antennae; head often with a Y- —Prothorax not hooded; antennae with a compact club shaped impressed line on vertex ... 38 ... 46
— 1 - 46(45) Form very convex, broadly oval; tibiae flattened, -Basal 3 ventrites fused, the sutures between them expanded towards apex; procoxal process received by shallower and narrower than those between 3rd/4th median cavity in mesosternum and 4th/5th ventrites (which are freely movable, ... (Fig. 172) .. [33] OSO E I AE usually with exposed intersegmental membranes); —Form less convex, elongate; tibiae not flattened nor antennae variable in shape, but very rarely with a 3- expanded apically; procoxal process absent segmented club; body length usually over 3.5 mm ... [34] E MES I AE (Dermestinae) ... (Fig. 219-221) [68] E E IO I AE
47(42) Tarsal formula 5-5-4 ... 48 56(54) Procoxal cavities closed behind by lateral expan- —Tarsal formula not 5-5-4 ... 69 sions of apex of prosternal process (Fig. 39); basal 3 or 4 ventrites fused together ... 57 48(47) Head sharply constricted to a narrow neck, best —Procoxal cavities closed behind by inward extensions seen in oblique views in most specimens ... 49 of hypopleura (Fig. 38), or not completely closed; —Head not sharply constricted to a narrow neck ... 53 ventrites usually free, rarely with basal 2 fused together ... 58 49(48) Apical abdominal tergite produced posteriorly into a stout spine ... (Fig. 213) .. [63] MO E I AE 57(56) Prosternal intercoxal process broad; basal 3 ven- —Apical abdominal tergite not produced as a spine ... 50 trites fused together, and 4th and 5th movable; scales usually present ... (Fig. 217) .. [66] U O I AE 50(49) Antennae flabellate; tarsal claws toothed (Fig. 87) (formerly as Zopheridae) ... (Fig. 214) .. [64] I I O I AE —Prosternal process very narrow between coxae; basal 4 —Antennae not flabellate; tarsal claws not toothed ... 51 ventrites fused together, only the 5th movable; scales usually absent ... (Fig. 218) .. [67] CΗΑ CO ΥΙ AE 51(50) Tibial spurs pubescent (Fig. 85) ... (Fig. 233) .. [75] SC A II AE 58(56) Tibial spurs serrate (exception: οxοz l r (Fig. —Tibial spurs absent ... 52 84) ... (Fig. 212) .. [62] ME A YI AE —Tibial spurs simple or absent ... 59 52(51) First 2 sternites fused, the suture between them obliterated, at least medially; tarsi with penultimate 59(58) Penultimate tarsal segment bilobed, extending be- segment small and simple, antepenultimate segment neath last segment ... 60 lobed below ... (Fig. 232) .. [74] A E I AE —All tarsal segments simple, without lobes ... 62 —First 2 sternites freely movable, the suture between them distinct; tarsi with penultimate segment lobed below, antepenultimate segment simple 60(59) Basal 2 ventrites fused together, and ventrites 3-5 ... (Fig. 228-231) .. [73] A ICI AE with visible intersegmental membranes ... (Fig. 229) .. [73] A ICI AE 53(48) Elytra abbreviated, leaving at least 4 sclerotised (Lagrioidinae) tergites exposed ... (Fig. 227) .. [72] SA I GI AE —Ventrites free, the lines between equally distinct... 61 (Inopeplinae) —Elytra not or only slightly abbreviated, covering 61(60) Elytral epipleura indistinct (Fig. 95), restricted to abdominal tergites, which except for pygidium are proximal (basal) half of elytra; body form as illustrated membranous ... 54 ... (Fig. 223) .. [70] OE EME I AE —Elytral epipleura distinct but narrow (Fig. 96), exten- 54(53) With a shelf-like canthus formed by dorsolateral ding to, or almost to apex of elytra; body form as margin of frons under which antennae are inserted, the illustrated ... (Fig. 224, 225) .. [71] Y OC OI AE canthus extending back and usually encroaching on front margin of eye (Fig. 11, f); antennal socket and 62(59) Body usually strongly depressed, usually narrow scape not visible from above ... 55 and parallel sided (Fig. 193-195) ... 63 —Without a true canthus extending back to eye; antennal —Body not strongly depressed, if broad then not par- socket and base of scape usually visible ... 56 allel sided ... 65
55(54) Only ventrites 1 and 2 fused, remaining ventrites 63(62) Body pubescent; tarsal segment 2 and/or 3 lobed movable, with visible intersegmental membranes; ant- below or, if not lobed, then antennal scape at least as ennae with a distinct, loosely articulated, 3-segmented long as the 3 following segments combined; pronotum club; body length less than 3.5 mm without sublateral carinae and often with anterior ... (Fig. 210) .. [60] A C EOC Y ICI AE angles produced... (Fig. 192, 193) .. [45] SI A I AE
— 1 9-1 9-- —Body appearing glabrous; tarsal segments 2 and 3 not -Anterior margin of pronotum almost always convex, lobed below; antennal scape as long as or shorter than covering head, which is not visible from above; if the 2 following segments combined; pronotum usually pronotum emarginate anteriorly, leaving head partly with sublateral carinae (Fig. 195) ... 64 visible (Orthoperus), then femoral lines absent ... (Fig. 207) .. [57] CO Y O I AE 64(63) Pronotum without 2 submarginal carinae ... (Fig. 194) .. [46] CUCU I AE 73(70) Body characteristically strongly depressed; head —Pronotum with 2 submarginal carinae sharply constricted to a distinct neck behind prom- ... (Fig. 195) .. [47] AEMO OEI AE inent temples; basal 2 ventrites fused ... (Fig. 222) .. [69] OS OMI AE 65(62) Procoxae strongly, obliquely projecting, contiguous —Body convex to moderately depressed; head without a or almost so, obscuring intercoxal process; body shape distinct neck; all ventrites free, or basal 3 ventrites as illustrated ... (Fig. 226, 227) .. [72] SA I GI AE fused ... 74 —Procoxae not projecting above level of intercoxal process, which is clearly visible between them; body 74(73) Trochanters of oblique clavicorn type (Fig. 60, d); shape not as above ... 66 terminal segment of maxillary palp more slender than penultimate segment, often needle-like; body form as 66(65) Elytra truncate, leaving pygidium exposed; anten- illustrated ... (Fig. 203) .. [54] CE Y O I AE nae apparently 10-segmented (segments 10 and 11 —Trochanters of heteromeroid type (Fig. 58, 59), with fused, forming a 1-segmented club in males) base of femur extending around trochanter on one side ... (Fig. 189) .. [43] MO O OMI AE (Monotoma) to coxa; terminal segment of maxillary palp broader —Elytra not truncate; pygidium not exposed; antennae than penultimate segment ... 75 11-segmented, with a 3-segmented club ... 67 75(74) Basal 3 ventrites fused, the lines between them 67(66) Form broad, convex, almost hemispherical; body fine and shallow in contrast with the broad, deep glossy, glabrous, 1.6-2 mm long (males of Cyclaxyra) sutures between ventrites 4 and 5 ... 76 ... (Fig. 196) .. [48] A AC I AE (in part) —Ventrites free, the divisions between them equal in —Form elongate, less convex; body matt to slightly depth and width ... 77 glossy, pubescent ... 68 76(75) Antennae 10-segmented; sides of head and ante- 68(67) Male mesocoxal cavities open laterally, reached rior part of prothorax with deep cavities for retracted by mesepimera; body form as illustrated; length 2.5-5 antennae (Fig. 8, b) ... [68] E E IO I AE mm (Fig. 190, 191) .. [44] OEOS IC I AE (Phrenapatinae) —Male mesocoxal cavities closed laterally by meeting of —Antennae 11-segmented; sides of head and prothorax mesosternum and metasternum, not reached by mes- without cavities ... [65] CO Y II AE (Pycnomerinae) epimera; body form as illustrated; length usually less than 2.5 mm ... (Fig. 198) .. [50] C Y O AGI AE 77(75) Antennae 9- or 10-segmented with a 3-segmented club (Fig. 9); head deflexed (Fig. 9); prothorax some- what hooded over head (Fig. 9); lateral margins of 69(47) Tarsal formula 4-4-4, rarely 3-4-4 in males frons explanate above antennal base (Fig. 9), but tan- (Mycetophagidae) ... 70 gential to top of eye, not forming a true canthus —Tarsal formula not as above ... 80 ... (Fig. 211) .. [61] CII AE —Antennae 11-segmented, usually with a 3-segmented 70(69) Tarsi with antepenultimate segment and often club; head not deflexed; prothorax sometimes extend- basal segment lobed below; femoral lines often pres- ing forward over head, but not hooded as in Ciidae; ent on ventrite 1 (Fig. 92, a) ... 71 lateral margin of frons often forming a canthus in line —Tarsal segments not lobed; femoral lines absent ... 73 with eye and often encroaching on front margin ... 78
71(70) Body form characteristic, with pronotum bearing 78(77) Mesocoxal cavities open laterally (Fig. 42), a conspicuous raised, curved, sharp carina on each reached by mesepimera; dorsal body surface smooth, side... (Fig. 204) .. [55] E OMYC I AE (Mycetaeinae) pubescent, without scales; male tarsi 3-4-4 —Body and pronotum not as above ... 72 ... (Fig. 209) .. [59] MYCE O AGI AE —Mesocoxal cavities closed laterally (Fig. 43) by meso- 72(71) Anterior margin of pronotum almost always sternum and metasternum, not reached by mesepimera;
emarginate, leaving most of head visible from above dorsal body surface often rough (e.g . , Fig. 215) and ... (Fig. 206) .. [56] COCCI E I AE clothed with scales; tarsi 4-4-4 in both sexes ... 79
-11 - 79(78) Body usually subcylindrical, glabrous or micro- smaller than segment 9, 10, or 11, or if antennae setose, glossy; antennal insertions exposed; all vent- filiform then elytra with transverse striae; protarsi rites free ... (Fig. 201, 202) .. [53] O I E I AE broader in males than in females; body moderately to —Body rarely subcylindrical, its surface usually rough strongly convex, oval in outline and clothed with hairs or scales, usually dull; antennal ... (Fig. 120, 121) .. [ 10] EIO I AE insertions concealed; ventrites 1-3 or 1-4 connate —Antennae not so; elytra without transverse striae; ... (Fig. 215, 216) .. [65] CO Y II AE protarsi usually not broader in males than in females ... 88
80(69) Tarsal formula 3-3-3 ... 81 88(87) Antennae weakly but distinctly clubbed, with —Tarsal formula 5-5-5 ... 84 segments 1-6 glabrous, segment 3 longer than scape, and segments 7-11 much broader, pubescent, forming 81(80) Body minute, less than 0.7 mm long; antennal a loose club; protarsi and mesotarsi expanded in males pedicel larger than scape; common in bat guano, and ... (Fig. 119) .. [9] AGY I AE known from nests of tt x l n —Antennae not so; protarsi and mesotarsi rarely ... [32] ACO SO II AE (Derolathrinae) expanded in males ... 89 —Body larger, exceeding 1 mm in length; antennal scape larger than pedicel ... 82 89(88) Body small, less than 2.5 mm long, glossy, very convex; elytra truncate apically, exposing sclerotised 82(81) Eyes absent; body parallel-sided pygidium; ventrite 1 at least as long as next 3 ventrites together ... (Fig. 131) .. [12] S A Y I I AE ... (Fig. 201) .. [53] O I E I AE (Anommatinae) —Eyes present; body not as above ... 83 (Scaphidiinae) —Body usually larger, if small then not glossy and strongly convex; elytra not truncate; pygidium not 83(82) Procoxal cavities open behind; body form char- sclerotised; ventrite 1 shorter than next 3 ventrites acteristic, as illustrated together ... 90 ... (Fig. 205) .. [55] E OMYC I AE (Holoparamecinae) —Procoxal cavities closed behind; body form not so 90(89) Body shape characteristically ant-like; head ... (Fig. 208) .. [58] CO ICA II AE (= Lathridiidae) strongly constricted to a narrow neck; prothorax and elytra narrowed at base, forming a distinct waist; 84(80) Metasternum longer than combined length of ven- antennae not clubbed; body strongly pubescent; trites 1-4; legs very short; body depressed, character- terminal segment of maxillary palp short and sharply istic in shape, as illustrated pointed ... (Fig. 122) .. [11] SCY MAE I AE ... (Fig. 171) .. [32] ACO SO II AE (in part) —Body shape not so; other characters not present in —Metasternum shorter than combined length of ven- combination ... 91 trites 1-4; legs longer; body shape not so ... 85 91(90) Abdomen with pygidium not exposed; antennae 85(84) Prothorax hooded (Fig. 10), produced over head; with last 2 segments fused, forming a conspicuous head partly or entirely concealed from above ... 86 club; body moderately to strongly elongate —Prothorax not hooded; head not concealed ... 87 ... (Fig. 188) .. [43] MO O OMI AE ( n x 86(85) Antennal insertions separated by more than length —If abdomen with pygidium exposed and sclerotised of scape; trochanters obliquely joined to femora (Fig. then antennal club 3-segmented and body less 60, a); elytra elongate, parallel-sided elongate ... 92 ... (Fig. 174-176) .. [35] OS IC I AE (incl. Lyctinae) 92(91) Antennae with a very broad, compact, 3-segmen- —Antennal insertions closely approximated on frons, ted club; pygidium, and sometimes 1 or 2 tergites in separated by much less than length of scape; troch- front of it, usually sclerotised and exposed anters squarely joined to femora (Fig. 61, d); elytra ... (Fig. 187) .. [42] I I U I AE broad, convex ... (Fig. 178) .. [36] ΑΝΟΒΙΙ ΑΕ —Antennae with club (if any) narrower and more loosely (Ptininae) articulated; abdominal tergites concealed under elytra ... 93 87(85) Antennae with last 5 segments distinctly broader than basal segments, forming a loosely articulated 93(92) Procoxae strongly projecting and contiguous club, with segment 8 smaller than segment 7 or 9 (Fig. externally, their cavities oblique; body with stout, 24), or rarely with a 4-segmented club with segment 8 erect pubescence (Fig. 182-184) ... 94 —Procoxae not or weakly projecting, not contiguous ex- -Elytra without erect setae on dorsal surface, without ternally, their cavities not oblique; body if pubescent definite rows of punctures; body shape not so; lateral dorsally then with pubescence shorter, finer, and less margins of pronotum and elytra usually not explanate projecting, or with short bristles ... 96 ... 103
94(93) Tarsi with at least segments 2 and 3 lobed below 102(101) Protibia with a single spur; antennal club (exception: M t x n all segments simple); antennae usually markedly asymmetrical usually with a loosely articulated, 3-segmented club ... (Fig. 180) .. [37] OGOSSI I AE (Lophocaterinae) ... (Fig. 183, 184) .. [39] C E I AE —Protibia with 2 unequal spurs; antennal club sym- —Tarsi with all segments simple; antennae filiform ... 95 metrical ... [37] OGOSSI I AE (Protopeltinae)
95(94) Body bearing very long, erect dorsal setae; mouth- 103(101) Body small, strongly convex, glossy, less than parts, especially mandibles, prominent; body shape 2.5 mm long; elytral epipleura each with 2 very deep, characteristic, as illustrated interconnected pits in females ... (Fig. 182) .. [38] C AE OSOMA I AE ... (Fig. 196) .. [48] A AC I AE (genus Cyclaxyra) —Not as above ... (Fig. 186) .. [41] ME Y I AE —Body not so, more than 2.5 mm long; elytral epipleura (also as Dasytidae) without deep pits ... 104
96(93) Antennae 10-segmented, with a strong, apparently 104(103) Body broadly oval; sides of pronotum explan- 1-segmented club formed by fusion of segments 10 ate; antennae with last 3 segments much broader than and 11; body shape characteristic, as illustrated the remainder in females ... (FIg. 185) .. [40] YCOSECI AE ... (Fig. 191) .. [44] OEOS IC I AE (Priasilphinae) —Antennae not as above, usually 11-segmented; body —Body elongate, parallel-sided; sides of pronotum not at shape not so ... 97 all explanate; antennae with last 3 segments scarcely differentiated from the remainder ... 105 97(96) Mesocoxal cavities open laterally (Fig. 44), i.e., mesepimeron reaching cavity ... 98 105(104) Prothorax with lateral gibbosities, but not car- laterally in females; body form as illustrated —Mesocoxal cavities closed laterally (Fig. 45) by inate OEOS IC I AE (Agapythinae) meeting of mesosternum and metasternum, which ... (Fig. 190) .. [47] —Prothorax without lateral gibbosities, carinate later- exclude mesepimeron from cavity ... 106 ally, at least at apex and base; body form as illustrated ... (Fig. 197) .. [49] CA OG A I AE 98(97) Body elongate, strongly depressed ... (Fig. 194) .. [46] CUCU I AE —Body usually less elongate, less depressed, and if 106(97) Elytral epipleura indistinct or well marked only elongate then strongly convex ... 99 in proximal half; epipleural carinae indistinct or absent posteriorly; body form as illustrated 99(98) Protibia with outer edge characteristic, carinate, ... (Fig. 198) .. [50] C Y O AGI AE (females) with regularly spaced narrow, transverse indentations —Elytral epipleura distinct, extending to apex or almost (Fig. 83); body usually very small and convex, less so; epipleural carinae distinct throughout ... 107 than 1.7 mm long ... (Fig. 179) .. [37] OGOSSI I AE (Rentoniinae) 107(106) Tarsal claws strongly toothed at base; body —Protibia with outer edge not so, not carinate; body broad, strongly convex length exceeding 1.7 mm ... 100 ... (Fig. 196) .. [48] A AC I AE —Tarsal claws simple; body less convex, more elongate 100(99) Procoxal cavities closed behind; elytra distinctly ... 108 striate ... (Fig. 181) .. [37] OGOSSI I AE (Trogossitinae) 108(107) Pronotum with distinct paired impressions —Procoxal cavities open behind; elytra without distinct basally; procoxal cavities open behind striae, though occasionally with numerous rows of ... (Fig. 199) .. [51] A GU II AE punctures ... 101 —Pronotum without paired impressions basally; pro- coxal cavities closed behind 101(100) Elytra with numerous long, erect setae on dorsal ... (Fig. 200) .. [52] E O Y I AE surface (Fig. 180), or with numerous rows of punc- tures; body broad, parallel-sided; lateral margins of pronotum and elytra explanate ... 102
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[ ] yso i ae a um i wi gs a se . ou u e a e a c es ig. 0 h z d t pr pr ( ou a a e ase o a s, ee i g o o e g ou we i g A ack, g ossy, su a a e si e ee e 6 mm o g, a o o s. I coas a a eas om o Waika o o , a i i g u e a k o i woo o a y ecaye ogs. K ow es ecia y commo o o s o e is a s. om o a , Auck a , Waika o, a G ea a ie . [ ] y isci ae: y o o i ae [2] Ca a i ae: Cici e i ae ig. 08 d pl t S a ig. 02 nd l t b r l t ( a icius A e o ga e o a , s ig y e esse , ack ee e 2. 2. A co s icuous y wo o e ige ee e 0 2 mm o g, mm o g. E y a s o g y u c a e, Iacki g s iae o a si mos y a k wi me a ic g ee e ec io s a i egu a a meso a si e a e i ma e, a me a a si wi a a e a c eam co ou e a eas o e y a. A e ae a egs i ge o swimmi g ai s. ima i y a o we e , o g a s e e eyes Ia ge a omi e e y a wi i a i i g a i e a s a 00 m as we as ow a a s iae, c ose y u c a e, a wi a ow o a ou 4 o eae em o a y o s, a e e wa m oo s i e e ma ea su u e ma i es Ia ge a s a oi e . A u a egio o o o ua, a ou i quie eac es o s eams. i y, o e a i a s suc as c ay a ks, w e e i eys o A u s a e ee o se e i ig . eco e om o o e i sec s. Mo es e y swi y, a akes o ig a i y. Ca e o S ewa I. a ae i e i u ows i e ose wa m a eas. Occu s o o mai is a s. [ ] y isci ae: y o o i ae g. 0 K h l dr phr t O is [2] Ca a i ae: Ca a i ae A sma , su e a ea i i g ee e . .6 mm o g, ig. 0 M r p b r f rb rn ookes a ow y e o ga e, su a a e si e , ow . i egs a A Ia ge, ack g ou ee e 20 24 mm o g, wi s ig y o y, a icu a y a e a y, wi o g, e ec , i e se ae i i esce u e o y si es. ea is i c y e o ga e, o o um wi si es a cua e. Co ec e om we s a o ec i g i o o o u i g emis e ica eyes ma o es i e o e a o e Sou I. [ dr n ft r i es s o g, oa . A e a o o s ai s. eco e mai y Ord h 6 ] om Wai oua o es , o a . [4] Gy i i ae [2] Ca a i ae: Miga o i ae ig. 0 G r n nv x l Mac eay ig. 04 x r n br d Gué i A ack, g ossy, oa s a e w i igig ee e 4. 4. mm A me ium si e g ou ee e mm o g, e is o g. A e ae s o , wi segme s co a a e i egs a k ow o ack wi e is ow a e ages. a e s a e , wi a i ge o o g swimmi g ai s. o o um a owe os e io y e y a o a . eco e om Occu s o e su ace o o s, owi g a i y o e e e Auck a Is. wa e a eyi g o o e i sec s. e i i e eye (a owe e a es i o see simu a eous y a o e a [2] Ca a i ae: Sca i i ae e ow e su ace. Mos ece y co ec e o e A i a a ig. 0 Cl v n b l C au oi a eau, o a . A mo e a e y sma , e u cu a e g ou ee e 6 8 mm Io g, a k ow o a mos ack wi e is a e ages [ ] y o i i ae: o e o i ae a asa a o e y a. A c a ac e is ic a ow wais ig. r l ph l r O c ymo e wee o a a e y a. a gi g om ce a o I. o A weak y co e , e o ga e ee e 2.0 .0 mm o g. o y S i i s ay i e o . ou i e i e u e e wee o o um a e y a o o a si es si ua e a e ae e y o g, wi a oose y o me [2] Ca a i ae: ec i ae segme e c u . esc i e om 2 s ecime s co ec e a g. 06 ll n l r ( ou ee o , a wi su seque eco s om Caw o A sma g ou ee e 4.0 . mm o g, ow wi a e , a k, e so , is u ique s ecies e ese s a e emic ye owis ow a e ages a a e a e y a. Eyes a ge ew ea a su ami y. S ecime s co ec e o ocks i os e io o o um s o g y a owe e y a wi 2 e s ay o e o wa e a s sugges a i may e os e o a e a o ec io s. Co ec e om u e ogs o e semiaqua ic. eac a om sa ea a s eam a Ka eka e a W a i u, ea Auck a . [ ] y o i i ae: y o i i ae ig. 2 En hr tr t ( ou [2] Ca a i ae: a a i ae A e o ga e o a , ye owis ow ee e 4.4 4. mm o g. g. 0 Ct n n th n v z l nd ai mai e Ma i a y a s o ge a a e ae e y a u c a e, o A e y agi e, e o ga e, g ossy ack g ou ee e 2 s ia e. A a e i e Aus a ia s ecies occu i g i ew mm o g. A e ae a egs i e y a egu a y s ia e, ea a i sma o s commo i ca e oug s i e use 2 omi e ac i e se ae a o e eye a a ow o Ke ma ec Is. A u s a a ae ee o a gae.
68 [ ] y o i i ae: S ae i ii ae [ ] Agy i ae ig. d t b l ou ig. ` r ph l pr l n t A s o g y co e , o a ee e 6. 8.0 mm o g, me a ic A mo e a e y Ia ge, oa y o a , e esse ee e .0 0 ack wi egs, ma i a y a s, a asa a e a mm o g, a k ow wi si es o o o um a e y a segme s a e a o e wi a e is i ge mo e a e y ye owis ow , e a a e. o a si a meso a si e o e se y u c a e. A e ae o g, wi a oose a e i ma e e y a s ia e a e ae wi a i is i c segme e c u . O igi a y esc i e om M A u , o segme e c u . eco e om e o I. wes e so we a e see s ecime s o esuma y is s ecies om . o o ua, Sa is u y, a M o e . [ 0] eio i ae: Camia i ae ig. 20 In t p l n ll ou [6] is e i ae: Sa i i ae A sma ca io ee e 2. 2. mm o g, e is wi ig. 4 h rdt p d tr x (S a w i is u esce ce. o o um a owe os e io y, wi a A o us , o o g, s o g y co e , g ossy ack i ee e a s e se asa im essio e y a o a i ou i e 4.8 .0 mm o g. E y a a e ia e , e osi g as seg a e ae wi a 4 segme e c u , i s 2 segme me s o a ome egs s ou , wi ume ous s i es o a owe a e o e s. ou i ea i e . eco e egs we a a e o u owi g. Occu s o sa y eac es, om e e so a ea. o e u e ea is o i s, occasio a y u e sea wee . eco e om o a o S ewa I. [ 0] eio i ae: Co o i ae ig. 2 C l n h rt l ( ou [6] is e i ae: Sa i i ae A o o g, mo e a e y co e sma ca io ee e 2. 2. ig. n l t p ( ou mm o g, u esce , u a k ow , wi egs a a A o a , co e , g ossy ack i ee e 4. 4. mm o g. e ae e a e a c u a ke , 4 segme e . e mi a A e ae s ou , ea i g a com ac c u e y a a e ia e segme o ma i a y a s sma a s a y oi e ea a ica y, ea i g ygi ium e ose o e egs a e e o wi a is i c eck. Wi e y is i u e , a eque y iggi g,wi a o ima e y 6 ee a o g ou e e ge, a caug i ig i e ce io a s. o e egs ea i g s ou is es. A Oma u a ou i a s o ai e a oos i a o ow a e kau i ee, o G ea [ ] Scy mae i ae: Scy mae i ae a ie Is a ou i ki g is e es s. A u s a a ae g. 22 Adr t l r ( a o a y ee o y a ae i gua o. eco e om A e o ga e o a ee e . . mm o g, e is ow , Oma u a S a e o es o Co is I. ea S ewa I. g ossy, s a se y u esce . A e ae 0 segme e a e o e y a o u y co e i g a ome as segme o [ ] y ae i ae: y ae i ae ma i a y a sma ea wi a eck, a a ma ke co ig. 6 d n l t p lp O is s ic io e wee o o a a a ome o mi g a is i c A sma casca e ee e . mm o g, e is ow wi wais . Occu s i ea i e . eco e mos y om e a e c y ea o e a ume a a ea o e y a. Ma i a y Sou I. (o igi a y ou i e ou e u a ey a om a s o g, eac i g e ea eyes e y a wi i e s ia e ce a o Is a ( ese oa . ea i g a ow ca i a. A u s a e ee co ec e om Oc o e o u e om s eams i e o I. [ 2] S a y i i ae: Mic osi i ae g. 2 M r lph l t r ou [ ] y ae i ae: Oc e i ae A sma ee e 2.2 2. mm o g, a k ow wi e is g. M r p th z l nd O is a si a asosu u a a o e y a. S a se y u esce A u i o m y a k ow casca e ee e 2. mm o g, wi e y a co e i g mos o a ome o i ae s o g, a a e , e se, s o g y ecum e se ae o o o um a e e , ea i g ee a s i es a e ae wi c u 4 e y a. ea a o a wi g a u ose mic oscu u e segme e . K ow o y om e eac a o C a me s, e y a g a ous e wee mic oscu u e a se ae. Occu s u e i , a oug mos o e s ecies i a i o es s a i i e , a is y ica y co e e wi e is amo gs e a e ee caug mai y i ig i e ce io a s. se ae. eco e om e C a am Is, u e i is ic , a S ewa I. [ 2] S a y i i ae: Omaii ae ig. 24 O l lb p nn (Kiese we e [8] i ii ae: i ii ae A me ium si e o e ee e . mm o g, mo e a e y ig. 8 t pt n d l n Ma ews co e a g ossy, wi ea ack, o o um e is A e o ga e o a , co e ea e wi ge ee e .0 . ow o ack, e y a e i e y ye ow o ack, a a e ae mm o g, ack wi ye ow egs a a e ae. o o a wi segme s ye owis ow , emai e a ke . wi si es s o g y cu e o o um i egu a y u c u e Occu s u e ecayi g seawee s o e s o e. eco e Occu s u e ecayi g seawee s o e s o e. eco e om e Auck a Is, Cam e I., a (e a imi a Mac om e Auck a Is, Cam e I., a (e a imi a Mac qua ie . [ dr n ft r St l 64.] qua ie I.
6 [ 2] S a y i i ae: o ei i ae [ 2] S a y i i ae: Sca i i ae ig. 2 S lph t l n t d ou ig. r t ll r ( e e ac e A sma ee e . .6 mm o g wi o y a e o a k A s o g y co e , ig y g ossy ee e 2.4 2. mm o g, e is ow , a some imes wi ye owis ume i a wi e y a o g, u ca e a ica y, e osi g sma , a e e os e io e y a a g es. ea wi a u c i o m o ea o ygi ium a e ae a egs a so a e e . ou u e ei e si e e y a co e i g ea y a o a ome . ou oose a k o ea u ks a a c es, u e s o es, o i amo gs ea mou . eco e om e Sou I. [ dr n mois ea i e . Some sca i ii e a ae a e associa e ft r St l 66.] wi u gi. eco e om e oo K ig s Is a s o Sou a . [ 2] S a y i i ae: se a i ae ig. 26 S l l n t ou [ 2] S a y i i ae: ies i ae A e o ga e, s e e , somew a e esse ee e 2.2 2. ig. 2 r n h d n (Came o mm o g, g ossy e , wi e y a, a e ae, a s, egs, a A su a a e , a e e ee e 6.0 8.0 mm o g, a k ow u esce ce ye owis . A is i c wais e wee o o a wi e is egs. ea a s e se, wi a e ae a o a e y a e y a ea i g segme s o a ome e ose , ima e y as o g as o y o o um wi a e io a g es wi ou s iae u wi c a ac e is ic os asa o eae. A o ec i g e y a g oo e . ou u e a k o ka aka is i c i e ami a o e u e si e o e ea , us e i (C r n rp l v t , a o a y sa o agous. e me um. Occu s i ea i e , es ecia y o se ges. eco e om e We i g o a ea. [ dr n ft r St l ai y commo i e Auck a a ea. 0 .]
[ 2] S a y i ae: oeoc a i ae [ 2] S a y i i ae: Oso ii ae ig. 2 d phl h r tr l ( au e ig. t t r h f rr n ( ou A sma ee e .8 2.0 mm o g, ye owis ow wi ea A e o ga e, a a e si e , e is ow ee e .4 . ow , o e i e y ow . A ome ea i g mes e mm o g. A ome cy i ica , wi e gi es a s e i es mic oscu u e. Occu s o o mai is a s, a may e use a e a y a a e gi es a se . ou i ea i e a a e i e i ew ea a si ce i is k ow a so om umus, mos eque y u e a ai e omi a e ege a Aus a ia a ew Ca e o ia. io i e o o e o Is a . eco e om o a o e so . [ 2] S a y i i ae: ac y o i ae ig. 28 S p d ph l p. [ 2] S a y i i ae: O y e i ae A sma , ig y co ou e ee e .8 2. mm o g, e is ig. 4 C rp l s . ow wi ea a os e io a o e y a a k ow , A sma , a e e , a k ow o ack ee e . . mm emai e o e y a ye owis , a a ome ye owis o g a ome oa e i g a ica y e ce o co ica os e io y a ow is asa y, ea i g o g o u i g a e . C rp l s ecies a e sa o ages occu i g i se ae. S p d ph l s ecies a a e a u esce o o es i e , u may e a ac e o u g a ca io . ey o um. ey a e commo i o es i e , o i g ogs, a a e o e a u a , u i e s u ie ee s ecies a e u gi a occu o o mai is a s. e e a e ume ous k ow o e a e i e. S p d ph l s ecies i ew ea a , se e a o em o ye ame . [ 2] S a y i i ae: Euaes e i ae ig. A n th t v n ( ou [ 2] S a y i i ae: a oce i ae A sma , e is ee e 2.8 .0 mm o g wi a a k g. 2 br r p ll r rn (G a e o s a s e se s o i a ica i o a ome o e o y g ossy A me ium si e , a e e ee e 2. .0 mm o g, ow a s a se y u esce o ake e y a e eme y s o wi o e o y s o g y g ossy a a ome mo e a e y a ome mo e a e y g ossy, wi e se w i is u es g ossy. is i c y i s i a e ae wi o g se ae a ce ce. ike a ew ea a euaes e i es i is e emic, o g, o u i g a omi a se ae. A a e i e s ecies, ig ess, a i a i s o es i e . eco e om e eco e om mi Ca e u y. o e a o e Sou Is a a e Waika o.
[ 2] S a y i i ae: A eoc a i ae [ 2] S a y i i ae: seu o si ae ig. 0 Al h r h nd Kl z & C os y ig. 6 d p rr e aue A o us , a ow y o a ee e .0 . mm o g, s o g y A sma , a e e , e is ow ee e 2.8 . mm o g. g ossy, ack wi a e ages a ce a a o e y a ea ea i g a s a ow e essio a e a o mi e e is ow . ike a s ecies o Al h r i as a is i c o o um wi cos ae e y a eac wi 2 o gi u i a u mi u e seu osegme e mi a i g e ma i a y a cos ae a 2 a e a o gi u i a ca i ae a ome wi a ia a s. Al h r occu s o o mai is a s a e s ig y c u e se ae. K ow om ea i e i th f Ke ma ecs is s ecies is es ic e o e Sou I. o es . d p s ecies occu mai y i o es i e a moss, a a ea o e e a o s. eco e om e ay o e y a e e so a ea.
0 [ 2] S a y i i ae: ae e i ae E y a eac wi o 4 oa , o gi u i a i ges egs ig. M d n z l nd ( e e ac e osso ia a e a c u segme s c u ky ma i es ia A me ium si e , su a a e , e is ee e 4. .0 mm gu a . e so e s ecies o a e emic ge us wi c ose Io g, o e wi ea a ase o a ome a ke . e e a i es i Aus a ia, ew Gui ea, a C i e. a ae i e i mi a segme o ma i a y a s mi u e. eco e om soi a o a y ea g ass oo s. A u s y u i g e o o , o I. ae e i ae occu i am a i a s i e e i g i s i g a summe . eco e om e Moko o es s o ea o ies o wa e , a a e e a o s a i au Is i e o o e omo i e sou e akes. em oy e o a iges io . [ 4] ogi ae [ 2] S a y i i ae: S a y i i ae ig. 44 r x b r ( i aeus ig. 8 C f l t r ou A me ium si e , e omi a y uscous ca cass ee e A Ia ge, g ossy ack ee e mm o g, wi some 6.4 6. mm o g, o a a co e , ugose. A e ae s o segme s o a e ae a a si e is , a u esce ce a s ou egs s e e , o o ious y mo i ie o iggi g. ye owis . A ome coa se y u c u e i iae co e e A a e i e Eu o ea s ecies eco e mos y i e wi u s i es. U ike mos C f s ecies, ea muc o e a o e o I., some imes a ig . a ge i ma e a i ema e. ou u e ecayi g sea wee , w e e o a u s a e a ge, a e ow a ae [ ] Sca a aei ae: A o ii ae a e e a o s o i o a am i o s. A u s y ea i y, ig. 4 A r d t n ( o e es ecia y i o su s i e, a a e ee co ec e om A s ou , g ossy ee e 0 mm o g, e is ow o Oc o e o Ma c . eco e wi e y om Ca e ei ga o ack, wi a e ages usua y a e . E y a eac wi u e i . im esse s iae o i iae eac wi a ge ee , a o e i iae eac wi a ge a sma e s i e [ ] uca i ae: Aesa i ae a e ae wi a segme e ame a e c u . Acci e a y ig. C r t n th p rr n Wes woo i o uce om sou eas e Aus a ia, w e e i is a A a ge, ow s ag ee e 4 20 mm o g, wi u is commo as u e es . a ae i e i u e s i e soi , ack o c es o o o um a e y a. A e ae o eme gi g o ee a ig . ey a e u usua i ee i g o e owe , wi a segme e se ose c u . a ae a e C g ass ea es a e a oo s, o e aggi g ea es i o s a e g u s occu i g i ecayi g woo o ka aka a e u e o co sum io . A u s y o ig o wa m o e ees. A u s a e some imes ou i e a a summe e e i gs. eco e om e o , es ecia y i a i a , a occu oug ou e yea . Wi e y is i u e , a eas wi ig o ca ic soi , a e Sou I. sou o M a some imes yi g o ig . G ey, Ca e u y.
[ ] uca i ae: uca i ae [ ] Sca a aei ae: A o ii ae ig. 40 r l t r t l t (Wes woo , ma e ig. 46 h h r n p ou A a ge, a k ow o a e y e is s ag ee e 2 20 mm A sma , s o g y co e ee e .0 4.0 mm o g, g ossy o g. ea a o o um wi u ye owis ow , e is ow o ack. egs wi oa , s i ose i iae a a esse sca es a isi g om coa se u c u es e y a s o a si mo i ie o iggi g ea g a u a e o a y, e icu a e , wi se e a a ow, e e a e , o squamose wi sma eyes e y a a mos g o ose, weak y s ia e . a s ma i es eac wi a s o g a ica oo , a sma ou i seawee o eac es. eco e om o mai su a ica o sa a e a oo , a a oa i u ca e is a s. asa o e. A u s a a ae a e commo y ou i ecayi g woo o e g ou , a a u s may a so e [ ] Sca a aei ae: Sca a aei ae ou u e a k o i i g ees, a icu a y th f . ig. 4 Onth ph r n l t o ema eco e om o mai is a s. A me ium si e ee e 8 mm o g, wi ea a o o um a ge y o e g ee a e y a a e u ous [ ] uca i ae: uca i ae mo e wi a k ow . E y a i e a s wi a ow o ig. 4 , 42 G d r r l t ( ou omi e g ossy g a u es, eac wi a e ec is e. A A Ia ge, g ossy, se ua y imo ic s ag ee e 20 2 mm a e i e Aus a ia s ecies o e associa e wi cow Io g. E y a wi e ec , ye owis ow sca es a isi g om a o se u g. K ow i ew ea a si ce e 8 0s, e se, coa se, ci cu a u c u es ea wi es e i a i g o a y a i e i s a e ma u e wi s ock eyes. Ma es a e e ea e a e a ma i es im o e y ea y se e s. Occu s o o mai is a s, i u ca e a e i s, wi a o sa oo ea e mi e. wi mos eco s om e Sou . eco e om e Waika o a Co oma e a eas. [ ] Sca a aei ae: Me o o i ae [ ] uca i ae: am imi ae ig. 48 Od ntr v n Wa ig. 4 ndr bl x rl W i e A a ge, ea s a e , u esce ee e mm o g A c oco a e ow , g ossy, ea i y u c a e s ag ee e ea e ow o iceous o o um a e y a e is 28 mm o g, wi co s icuous u ous ai s e a y. mo e wi a k ow scu e um a k ow e a
— 171 — su ace ac eous o co ae a s e a. A e ae i o e ose . Occu s i coas a a eas, o sa y eac es se es ame a e. As i a c a e s, e egs a e a a e o a o e ig wa e ma k, w e e a ae may e ou u e iggi g. A u s a e ou o ea es o s u s a ig , a ogs. eco e om e o Is a a e e so a ea some imes y o a i icia ig . a ae o mos s ecies o o e Sou I. Od ntr occu i o es soi , a occasio a y i soi o a i e ussock g ass a .O e o a g ou o a i e o es [ 6] Sci i ae o ms a gi g oug ou e Sou I. a o o e a ges ig. A pl t p p ll rn ou o e o I. A sma , e o ga e o a , mo e a e y u esce , e is ow ee e . 2.0 mm o g, wi a e ae a egs [ ] Sca a aei ae: Me o o i ae ye owis . e a su ace wi o a e essio s i o w ic ig. 4 C t l tr z l nd W i e ea , a e ae, a egs ca e o e a e ae wi a A mo e a e y a ge ee e 0 mm o g, ow is ack o egu a 6 segme e c u , a wi o ima segme s a e ow , wi e ge o c y eus a eyes a ke i egu a i o m egs o i g oge e ike e a es o a a e ae a (usua y e y a es aceous o o um a e ocke k i e. A u s a e ee ea e om o iage o e e is ow o ow ack, a scu e um o e e is ee e n rr a e o us i ge ea e a su ace a e ow . ygi ium a y e ose , e so . O igi a y ou o M i o gia a si ce co ec e smoo a g a ous e ce o ma gi a is es. Com a Wai oua o es , u ua a ge, M A ow a a (Gis mo y k ow as e g ass g u , a ame mo e o e y o e , awso a s (M Egmo , a akaka ( e so . a ie o e a ae, w ic i e i soi ee i g o e oo s o g asses. Ma u e a ae u a e i e soi , usua y i a e [ 6] Sci i ae Augus o a e Oc o e , a e ee es eme ge i o ig. 4 r n t tr t ll W i e em e . O wa m, s i ig s ey y, a a ig o e A mo e a e y a ge, a ow y o a , s ig y e esse ea es o a ious oo a s suc as u i , um, ose, ee e .0 0 mm o g, a e ow wi a e ae a egs a e, s aw e y, a Ar l p n . A e ma i g, usua y ye owis ow . E y a wi 6 cu e o gi u i a cos ae o e oo a s, ema es ay ei eggs i mois soi . u e su ace wi i e, ecum e u esce ce ea a ae a c i 6 2 ays a ass oug ee i s a s usua y s o g y e ow wa s e u ima e a sa as i g om o em e o e o owi g Se em e . E segme i o e . Co ec e y ea i g s u s. Wi e y emic o ew ea a , om o Ca e o u , u o is i u e o o mai is a s. commo i o a a Auck a , a o e a e i a i e ussock g ass a s. Causes mos amage i e [ ] Euci e i ae Sou I., es ecia y i O ago a Sou a . ig. E n t t rt ( ou A sma , oa y o a , g ossy ack ee e . .6 mm o g. [ ] Sca a aei ae: Me o o i ae ea a o o um s a se y u c a e e y a wi c oss ig. 0 r d ntr l ou s ia io s a e ae cy i ica , segme e , e as A a ge, a e e is ow ee e 6 mm o g segme ack me aco ae a ge, use o mesos e um a e ae 4 ame a e i ma e a ame a e i ema e egs s e e , e i egs a ges , wi sma s i es. A u s c y eus a o s e se y a coa se y u c a e o a a ae o a y ee o s ime mou s. isco e e a o um s o g y co e e y a s ia e, s o g y co e , oa ma s ea ee o , a si ce ou a S a o ike y u ca e e i , wi s ou e s ea i g a os e o a e a o o e wi e y is i u e w e e e s ime mou s occu . e essio . K ow o y om oug as u e a i a e s ic e a ea o C omwe , Ce a O ago. [ 8] C am i ae ig. 6 Cl b d t ou [ ] Sca a aei ae: y as i ae A mi u e, oa y o a ee e .0 mm o g, a e o a k ig. t r n h r t r ( a icius ow , s o g y g ossy, s a se y u esce . ea e y A mo e a e y a ge, s ou , co e , g a ous ee e oa , e e e e ea o o um a e ae 8 seg mm Io g, s o g y g ossy ack. Mi e a i emo a me e , wi a 2 segme e se ose c u egs s e e , wi eac wi 2 s o g s u s. Commo y k ow as e ack i iae Iacki g s u s. A a e i e Aus a ia s ecies, u y ee e, is s ecies was acci e a y i o uce om Sou wi ge , a commo y occu i g i aw c i i gs, ga e A ica. A u s some imes y o ig a ig . a ae ee o u i gs, com os , a i e i a i e o es . e oo s o g asses, a may cause su s a ia amage o aw s a as u es. Wi e y is i u e i wa me a eas o [ ] u es i ae e o I. ig. d n S a A mo e a e y a ge, somew a e esse , u e s a e [ ] Sca a aei ae: y as i ae ee e 8 mm o g, me a ic g ee wi 2 o ique ye ow ig. 2 r pt tr n t a icius s o s ma es wi o s e is . E y a oi e a ica y. A a ge, s ou ee e 2 0 mm o g, a k ow , o e Commo y k ow as e eec u es i , e u y wi ge c oco a e ow me ia y. egs oa , a a e o iggi g a u s ee i summe o ea es o th f , a a e a o o um i ma es ea i g a u a ica o ygi ium o e ia es o eec o es s.
— 2— [20] y i ae: y i ae [2 ] i o ac y i ae: A c y a si ae ig. 8 h r h tt n ascoe ig. 6 rr r pt r h rt ou A mo e a e y sma , oa y o a , co e ee e .6 . A mo e a e y a ge, c ose y u esce ee e 6 mm o g, mm o g, s i i g ack wi g ee is e ec io s. A e ae wi o y me ium ow a egs a a e ae ye owis . i se e o si es o o s, wi a segme e c u e y a A e ae se a e scu e um ea s a e i ma gi o eac wi a ou o gi u i a ows o u co ec e coa se o o um wi u ee o mi g a a s e se com . u c u es a omi a s e i es e i me aco ae wi a A u s a e ee co ec e e wee Oc o e a e ua y. a ge o a e essio o ece io o e ac e i egs. a ae i e i sa ea e wa e s e ge i i e e s. A u s a e o e ou u e s o es su ou e y moss, eco e om Auck a o mi Ca e u y. usua y associa e wi a ae, a a e ee o se e ee i g e a i e aisies (C l . eco e i e [26] C e o a ii ae sou e Sou I. om e G am ia Mou ai s (Mac ig. 64a, r n th r S a ke ie asi o e g ea ock mou ai s o Ce a A mo e a e y Ia ge, a ow y o a , s ig y g ossy ee e O ago, a a i u es om 0 m o o e 000 m. 6 mm o g, ack wi a si ye owis e a e y a io e o u e i ge mo e a e y e se y u c a e a u es [2 ] yo i ae ce . ea a ia y co cea e om a o e a e ae wi ig. rn d r t ou segme s 6 0 eac ea i g a o g, i wa y o uce A mo e a e y sma , mo e a e y g ossy ack ee e . o e o o um um s a e i a e a iew. e o y 4.0 mm Io g, co e e wi i egu a ye ow ai s. A e ae k ow s ecime s a e e o igi a wo co ec e y ou s o , wi a c a ac e is ic asymme ica c u o o um ( oca i y u s eci ie a a o e Ma aise a e a Eas wi a ee y im esse g oo e o ei e si e ea a e a Ca e. ma gi . O e o a um e o e es ia yo i ae ( e yo i s i o e oogeog a ica egio s a e mos y [2 ] Euc emi ae: Me asi ae aqua ic . A u s a a ae occu commo y oge e i ea ig. 6 h r pl x S a i e o th f o es s. A mo e a e y sma , e o ga e o a ee e .8 4.0 mm o g, ack wi g eyis u esce ce. A e ae o g, e 2 a [22] E mi ae: a ai ae segme s sma es e y a wi ou e i i e s iae. ou ig. 60 d r p ou o o es g ow a i ea i e a woo mou , a as A mo e a e y sma , oa , s ig y e esse , ow is ee ake i Ma aise a s. eco e om Kai u i ack ee e .4 . mm o g. ea a ow, wi omi e o a o um i ge i Sou a . eyes a e ae wi ou a is i c c u os e io e ges o o o um s a e y a wi ows o u c u es o y [28] E a e i ae: Ag y i ae c o e wi ai y s o u esce ce. Wi e y is i u e n ig. 66 h r f ld S a i e e s, o s o es a e wa e s e ge. A a ge, a ow y o a , ackis ow c ick ee e 0 mm o g. Si es o o o um e e y cu e a e io y o [2 ] im ic i ae: im ic i ae u os e io a g es e y a eac wi a ou s iae ig. 6 n h n r p ou a e ae s e e egs s ou me aco ae e ca a e A sma , o a , co e , g ossy ack ee e 2. 2. mm o g, os e io y o ecei e e ac e emu . O e o ou Ia ges c o e wi ye owis sca e ike se ae a a ge i wa y c ick ee es, qui e a ia e i s a e a si e. a ae a e a e s, e se ae i e a ai ike o o o um. A e ou i ea woo i o es s, w e e ey ey o a ae o ae wi a sma 2 segme e c u , i se e o ase o o e ee es suc as ce am yci s. A u s o is wi e y o s. Occu s u e s o es i s eams a a e ma gi o is i u e s ecies y ea i y o ig . i e s. A so ou i e es ia a i a s i mois i e a u e s o es, u ima i y a aqua ic s ecies. eco e [28] E a e i ae: e co i ae a ow u , us o o C omwe , i e Mo u ., ay o ig. 6 A h nd z ascoe e y, a i e Ke eu S m, Eas Ca e. A Ia ge, ow is c ick ee e 6 2 mm o g, a iega e a a ia e i co ou , esem i g a k. o o um oa e [24] e e oce i ae: e e oce i ae a o g, ou e a si es, a wi omi e os e io ig. 62 t r r n v l nd C a e ie a g es e y a a owe a o o um, a e i g os e io y A mo e a e y sma , e o ga e, co e , ow is ee e o y su ace mo e a e y oug . Occu s i c e ices i ocks . 4.0 mm Io g, ea i g ume ous is es. A e ae a u e ogs a s o es. eco e om e is a s o s o a ick e y a wi o a , ye owis ow s o s e Cook S ai a e C a ams, is e emic ew i iae a e e a s i ose, ackis ow . e o y ea a s ecies is co si e e o e e a ge e . s ecies o e e oce i k ow om ew ea a , om a ew wi e y sca e e eco s. [2 ] yci ae ig. 68 rr t r f p nn ( a icius A mo e a e y a ge, so o ie , e o ga e, e esse ee e 0 mm o g, ack wi e y a o a ge (o e ig
— — e i i i g s ecime s . A e ae ee y se a e e y a ma es wi a c u co sis i g o o g a c es, a i wi i e i i e o gi u i a i ges a a ge u c u es ema e sim e, wi c u segme s sma e a s ig y e wee em. eco e om e o I., es ecia y asymme ica . esc i e o e asis o a ma e a a o e a eas, a a ou e so i e Sou I. ema e ou ea W a ga ei ea s a a ai ua.
[ 0] Ca a i ae: ysmo oce i ae [ ] os ic i ae: i o e i ae ig. 6 A l f lv th r x ou ig. n d r n t ( a icius A me um si e , so o ie , ai y a so ie ee e 4.8 A sma ee e 2.0 .0 mm o g, a k ow wi e e y a. .0 mm o g, ack wi o o um ig o a ge. A e ae ea wi o s e se y u c a e o o um coa se y s e e a s ig y se a e, i se e o o s us e o e as e a e a e io y, e se y u c a e os e io y e y a eyes. A u s may e ea e om o iage o a ious ees i egu a y u c a e wi s o , e ec se ae. A a o ica a s u s, es ecia y w e i owe . Commo i e es s ecies e e o i g i am oo, a some imes i o I. o o Wai omo, i c u i g e e a C icke s mai e. eco e om Auck a . Is a i e a ie . [ ] os ic i ae: yc i ae [ ] e o o i ae: a ico ii ae ig. 6 t br nn S e e s ig. 0 th d r d nt rl C owso A sma o mo e a e y a ge, ow ee e 2.0 .0 mm o g. A sma , oa y o a ee e . 2.0 mm o g, somew a U e su ace wi ow ecum e u esce ce ea e esse , ow is o ack. ea wi c a ac e is ic a o o um wi coa se, s a ow u c u es e y a e essio s a wi 2 oce i, eac e o ma gi o eye s ia e, wi i e u c a io o o um s ig y e esse eyes omi e , coa se y ace e a e ae wi a me ia y os e io ma gi o 4 e i e wi i e u es segme e c u e y a wi ee s iae. A u s a e ee ce ce i o se es. A cosmo o i a es i a wi e a ie y co ec e om a ua y o Ma c . isco e e a A u s o im e s. eco e om o mai is a s. ass, a e o e om se e a oca i ies e wee e e so a ea a Ca es o , ea ee o . [ 6] A o ii ae: A o ii ae ig. dr br n ( um e o [ 2] aco so i ae A me ium si e , e is ow ee e .0 8.0 mm o g. ig. S ph ph n t S a ea co cea e y o o a a e ae wi a A sma , e o ga e o a , e esse , e is ow ee e segme e c u co sis i g o e o ga e a asymme ica . 2.0 mm o g. o o um a o ima e y ea s a e , segme s o o a co s ic e os e io y ea ase wi a o ou ce me ia e essio a wi coa se e y a wi ows o u c u es mos o o y su ace wi u c u es ea ess coa se y u c a e a e ae s aig , s o u esce ce. A u s a a ae occu i o , a y wi a oose segme e c u e y a wi ou is i c ecaye mu ogs a i imu oo i g a o e im e s s iae, u wi u c u es a a ge i somew a i egu a , ey usua y o e i o we im e . Wi e y is i u e , om cu e , o gi u i a ows egs a e s o a s ou . o a o e aie i ai s, u e i . Co ec e i e e so a ea, a ic o , a ea M Cook. [ 6] A o ii ae: i i ae [ ] oso e i ae ig. 8 t n p ou ig. 2 d ndr n z l nd S a A sma , oa y o a , s i e ike ee e 2. 2.4 mm o g, A mo e a e y sma , o a , s o g y co e , g ossy ack wi a ee ow a w i e s i e ac oss e y a a ee e 4. 4. mm o g, i e y a c ose y u c a e. E y a a i io a w i e ma ki gs e i s ou e s o y c o e wi ows o a ge u c u es. Wi e y is i u e i e o wi o g ye ow ai s. ea usua y s o g y e e e a I., a a so eco e om e Sou I. a ia y co cea e om a o e a e ae wi ou a c u egs a a e ae o g a s e e e y a muc oa e [ 4] e mes i ae: Mega omi ae a o o a . A u s occu i ea i e . Wi e y is i u e ig. r d r t ou i e o I. a o e Sou I. A sma , oa y o a , s ig y e esse ee e 2. .0 mm Io g, ack, co e e wi ack a w i is ecum e ai s [ ] ogossi i ae: e o ii ae o mi g a c a ac e is ic a e . A e ae o ma e u y ig. nt n d t v ntr C owso se a e, ose o ema e weak y c u e . A u s may e A mi u e, o a , s o g y co e , a e e is ow ee e ou o owe s, a a ae o ea e s a ea i s. . mm o g. A e ae a egs s e e ea oa eco e om e o o e o I., as a sou as u e su ace wi i e u esce ce e y a wi ou s iae Okauia ea Ma ama a. isi e s e i e wi a me ia kee . isco e e i ma e owe s o " n n n " ( o a y i e o o . r d t i [ ] os ic i ae: Eu e i ae Wai oua S a e o es , o a . A so co ec e i o e ig. 4 E d r ou o I. oca i ies i o es ea i e , a occu i g as a A mo e a e y sma , e o ga e ee e 4.0 .0 mm o g, sou as e so . ow , co e e wi w i e a ow sca es. A e ae i
— 4— [ ] ogossi i ae: o oca e i ae co ou e , mos y u ack wi e y a s ou e s o e i e ig. 80 Gr n r l r S a e y a a e ye owis , o e y a a ea i g ye owis wi A me ium si e , mo e a e y co e , e a y a ee e a ious com i a io s o ow o ack s o s. Occu s o . .4 mm o g, e o ga e o a wi si es mos y a a e . sa y eac es. eco e om o mai is a s, e Co ou e omi a y ack e y a wi w i e ma ks C a ams, a e Ke ma ecs. a iega e wi a e e , coa se y u c a e u o s ia e Io g, w i is ai s co e i g mos o o sa su ace [4 ] Me y i ae: asy i ae ma i a y a s omi e , e as segme a c e ig. 86 ` t b n ou s a e . Co ec e y ea i g o es o iage. a ae a e A me ium si e , so o ie , e o ga e ee e .8 4.0 mm some imes ou u e oose a k, w e e ey o a y o g, s ig y e esse , me a ic ue. ea wi ou a ee o o e i sec s a u s oo a e o a y e acious. is i c eck eyes omi e a e ae i i o m, wi ou a Wi e y is i u e . c u e y a u esce , wi ou s iae. Commo o owe i g s u s, a w e a e e akes o ig a i y. a ae [ ] ogossi i ae: ogossi i ae i e u e oose a k o ea a c es. Occu s o o ig. 8 p d pt r x n r p r (W i e mai is a s, u wi mos eco s om e Sou I. A a ge, e o ga e, a a e si e , s ig y co e ee e 2 mm o g, g ossy g ee is ack. ea a o o um [42] i i u i ae: i i u i ae wi e esse a eas ea i g ye owis w i e sca es ig. 8 l t p d p r ll ou e y a wi 4 o gi u i a kee s. A u s a e ee ou A me ium si e , oa y o a , s ig y e esse ee e u e a k a i e s ea o a a e ikau a m ea . 4.8 .2 mm o g, mos y a k ow , wi a e i egu a Co ec e om Se em e o A i . Wi e y is i u e , wi ma ki gs. Eyes s o g y co e ea , o o um, a eco s a gi g om i e a ie I. o Si e s eam ea e y a i e y u c a e e y a wi ou s iae ma e wi i We i g o . emo a e a e owa s a e . A u s a e ou o ee u ks, es ecia y wi owi g sa , a some imes i e [ 8] C ae osoma i ae ase o a e ea ikau a m ea es. Occu s e o mai ig. 82 Ch t r t d Wes woo is a s, u wi mos eco s om e Sou I. A Ia ge, e o ga e, su a a e si e ee e 4 mm o g, coa se y u c a e, ack wi a e e s ou e s a e y a [4 ] Mo o omi ae: Mo o omi ae a e . o sa su ace wi o g, e ec ai s a e ae a ig. 88 n x r nd S a egs s e e o o um su qua a e, wi a s o g y g ossy A me ium si e , a ow y e o ga e, su a a e ee e 4.0 me ia e e a io . Co ec e om u e a k o ogs a 6.0 mm o g, usua y a k ow o ea y ack, wi s a i g ea ees o a ious s ecies. Wi e y is i u e . a e ages a e , usua y us ow g ossy, wi mi u e, s a se a i co s icuous u esce ce. ea s o , as [ ] C e i ae: a oc e i ae oa as o a , wi 2 ee ca i ies e i eac eye ig. 8 M t x n rn t ou a um co cea e o o um a ow y e o ga e, wi o A me ium si e , s ig y a e e ee e 4.0 6.0 mm o g, a g es s a a s ig y o uce , a e a ca i ae s a , ow wi w i is ye ow, si ua e, i egu a y s a e e y a a wi 2 ows o Ia ge, ee u c u es i me ia a o s o s. o sa su ace wi o g, e ec s a se se ae ea isc e y a a ow y e o ga e, wi 8 u c a e g oo es. a o o um wi o gi u i a y e o ga e u c u es, a Co ec e u e a k i eec ( th f o es s, a e y a wi ou o es. O igi a y esc i e om oke some imes y ea i g o es g ow . Wi e y is i u e , i e , Ca e u y. We a e see a i io a s ec me s es ecia y i e Sou I. om u Mou ai , ea e so , a a u e e mi e s ecime o e same ge us om o a . [4 ] Mo o omi ae: Mo o omi ae ig. 8 M n t p n ll Au é [ ] C e i ae: E o ii ae A sma ee e 2. 2.2 mm o g, e i e y us ow , o a k ig. 84 h t ph v l ( a icius ow wi us ow e ges a a e ages. ea a A me ium si e , e o ga e o a , s ig y e esse ee e o o um wi coa se, oa , e se u c a io ea 6. .0 mm o g, mai y io e , wi is i c ye ow ma k a u y co s ic e e i , wi o u i g eyes o i gs o e y a. Eyes omi e a e ae wi a e o ga e o um wi a e io a g es o uce a a e a ma gi a asymme ica segme e c u 2 a sa segme s se a e e y a s o e e , wi ows o u c u es u es s o g y o e e y a coa se y u c a e, wi ou s iae, ce ce s o , i co s icuous, ye owis . A u s o is i o c o e wi o g, o u i g ai s. A u s occu i summe uce Eu o ea s ecies a e commo i aw c i i gs, o owe i g ees a s u s, w e e ey o a y ey o ga e i e , a com os , i a ocks w e e s ee o e i sec s. s e e , a i c icke s aw. eco e om Auck a .
[40] ycoseci ae [44] oeos ic i ae: Aga y i ae ig. 8 h l b t ( a ic us ig. 0 A p th f v ll ou A sma , e o ga e o a ee e 2. .0 mm o g, a ia y A mo e a e y sma , a ow y e o ga e ee e .0 .2 mm
— — o g, us ow wi a k ow s o s a 2 a e, [4 ] aemo oei ae a s e se s o s o e y a. Mo e a e y g ossy, s a se y ig. M r br nt l n t ( ou u esce , wi e essio s o o o um a e y a a A sma , a e e ee e 2.8 .0 mm Io g, ow wi a wi i co s icuous u e c es a e ae ea ike, wi a e is i ge, s ig y g ossy, wi i co s icuous s o weak y e i e segme e c u . A u s a e ee u esce ce. ea e o ga e, wi a me ia i ge a 2 co ec e om ecem e o e ua y. O igi a y ake a a e a i ges a e ae wi sca e e a ge a asym ou e u , u a so k ow om e o y o y, a oo me ica , e ice oa e e a ica y, a emai i g seg u (M A u a e a , e so , akaka i , M o e me s s o g y e o ga e o o um oa es a ica y, wi ( e so akes . . , u e Wai oa y, a Maka o a. 2 su a e a i ges e y a wi 0 o gi u i a i ges. K ow om e o I. a ee Ki gs Is. [44] oeos ic i ae: iasi i ae ig. r lph b r ou [48] a ac i ae: Cyc a y i ae A me ium si e , oa y o a , a e e ee e 4. .0 mm ig. 6 C l x r pr ou Io g, a owe os e io y, ackis ow , coa se y A sma , s o g y co e ee e 2.0 2.2 mm Io g, g ossy u c a e. U e su ace wi g eyis u esce ce g ou e ack wi egs a a e ae e is c es u . ea a i c um s o e y a e y a wi ou s iae, s ig y e a a e e y a s a se y u c a e a e ae wi a segme e a e ae wi a is i c segme e c u a e ae a c u e y a o s ia e. esuma y u gi o ous, occu i g egs s e e . A u s a a ae occu i ea i e o eec i soo y mou s o th f a Ol r . K ow om ( th f o es s. Wi e y is i u e o o mai o mai is a s. is a s. [4 ] Ca og a i ae [4 ] Si a i ae: o i ae ig. n d l h th n Wa ig. 2 r nt pr pl r l ou A sma , e o ga e o a , mo e a e y e esse ee e 2. A mo e a e y Ia ge, a ow y o a , a e e , ow ee e .2 mm Io g, a k ow wi egs a a e ae e is . 8.0 mm Io g. ea s ig y a owe a o o um ow su ace e a i e y u , wi i e mic oscu u e. a e ae wi sca e e eme y e o ga e, as Io g as o 4 ea e se y u c a e a e ae wi a oose seg o owi g segme s com i e o o um wi ee i e me e c u o o um wi si es weak y a gu a e us a io s o a e a ma gi s e y a o a a a a e a y, wi e i mi e e y a e o ga e, wi es a mi e g wi gs o gi u i a ows o u c u es o sa su ace some imes es igia scu e um a s e se egs s ou , wi o a si a ea i g i egu a ieces o a ma e ia a e i g o sa y meso a si (segme s 4 e a e i ma e. Occu s i sec e io . A u s a a ae occu u e a k o ea ee i s es s o e C a am Is. u ks a a c es o a i e ees. Commo i mos a s o e o I., wi a ew eco s om e Sou I. [ 0] C y o agi ae: C y o agi ae ig. 8 h rt v l ou Si a i ae:[4 ] Si a i ae A sma , o a , co e ee e . .8 mm Io g, e is ig. Or z ph l r n n ( i aeus ow wi s ou e s o e y a a a c es ea e y a A sma , a ow y e o ga e ee e ow 2.8 .2 mm Io g, a e a e mos o u e su ace wi s e e se ae. o e wi us ow a e ages o sa su ace co e e A e ae wi a oose segme e c u , ei i se io s wi s o , ye ow, ecum e ai s. ea e o ga e, wi e esse i o a e e a e su ace a se a a e y a em es as o g as iame e o eye a e ae wi a o gi u i a ca i a a e a ca i ae o o o a wi ou segme e c u o o um wi 6 ee o ei e si e se a io e y a wi i e u c a io a wi ou s iae e y a wi o gi u i a ows o u c u es. A cosmo o i a wi gs a se . Occu s i ea i e , es ecia y i th f es o s o e o uc s, a e i e i ew ea a . A u s o es s. isco e e a oa ma s ea ee o , a a e a a ae i es ce ea s, ie ui s, a oi see s. K ow ou o S e e s I. a mo e wi e y o e wes e si e om o mai is a s. o e Sou I.
[46] Cucu i ae [ ] a gu ii ae: e osce i ae ig. 4 l t p. ig. p l p pr l x (S a A a ge, a ee e 20 mm o g, a k ow , wi ea A me ium si e , a ow y e o ga e ee e .0 6.0 mm a o o um ea y ack, a e y a u i o m y a k ow Io g, su a a e o s ig y co e , ye owis ow . o e is a ica y, coa se y u c a e. ea a s e se, A e ae wi a e o ga e segme e c u o o um os e io y s ig y oa e a o o um ma i es wi ou a asa im essio o g oo e e y a wi ows o s o g, cu e , wi i e ee a e ae mo i i o m, wi u c u es. A u s a a ae occu o ea ea issue o asa a segme s s o g y e o ga e a 2 ee e s a ikau a m. K ow om e o I. a e segme e uce a ea s a e o o um wi se a e o e a o e Sou I. a e a ma gi s. A e emic ew ea a s ecies k ow o y om e ee Ki gs Is.
- 6- [ 2] E o y i ae: ac i ae [ ] E omyc i ae: o o a ameci ae ig. 200 h ll p l t W i e ig. 20 l p r t n ei e A me ium si e , e o ga e o a , co e ee e .8 6.0 mm A sma , e o ga e o a , e is ow ee e . . mm o g, ackis ow , s o g y g ossy. ea wi omi e Io g. A e ae wi a Ia ge, 2 segme e c u o o um eyes a wo ee g oo es a e ae wi a oose co e , es ic e e i o a is i c wais , w a se ies segme e c u e y a wi i egu a o gi u i a ows o o a e a y e o ga e o eae o mi g a asa im essio i e u c u es. A u s a e ou o acke u gi a e y a u esce , wi ou s iae egs s e e , wi i ia occasio a y i ea i e o u e a k. Wi e y is i u e i s u s. Associa e wi ea i e u e o i g ogs a wi e o I. u gi. ai y commo , a o a y wi e y is i u e .
[ ] o i e ae: A omma i ae [ 6] Cocci e i ae: Cocci e i ae ig. 20 An t d d tr t ( Mü e ig. 206 C n ll l n n a icius A sma , su a a e ee e 2.0 2.2 mm o g, o sa y A me ium si e , emis e ica a y i . . mm o g, g ossy, e igme e , a e ow , somew a a e e , wi wi ma y co s icuous, egu a o a ge ma ki gs o a s o , s a se a i co s icuous se ae. ea a ia y ack ackg ou . A u s a a ae ee mos y o a i s co cea e y o o um, is i c y a owe a a ica occu i g o g asses a ussock, u a e k ow om a o o a ma gi eyes e uce a e ae 0 segme e , wi e a ge o a s. eco e wi e y o e o mai wi a Ia ge segme e c u o o um a e o a , wi is a s a om sma e is a s c ose o s o e, om ea a ge, o gi u i a y e o ga e u c a io e y a wi a ow sea e e o su a i e. e i eu a, a wi 6 ows o se ose u c u es o ei e si e. A u s eco e om o e woo u ie i ea , [ ] Co y o i ae: Se ico e i ae u e ee y u ie s o es, a om ee i a a s ig. 20 An r t h rp Ma ews sea e o a e o . A i o uce y ogea Eu o ea A sma , e se y u esce , g ossy, ig o a ge ee e s ecies, eco e om y ie , Auck a . .2 .4 mm Io g. ea co cea e y o o o o um a e ae 0 segme e , wi a segme e c u o [ ] o i e i ae: o i e i ae o um wi i a g es s o g y o uce os e io y ig. 202 A t d r b l t ( ou e y a wi ou s iae, s ig y u ca e, e osi g a o A me ium si e , e o ga e o a , s ig y e esse ee e ygi ium 2 a sa segme i o e . ou o a u e .0 6.0 mm o g, a k ow o ack wi mos o su ace ea i e a e ase o se ges a a , a i coas a c i coa se y u c a e. A e ae i se e i o o eyes, wi ege a io a ou Auck a . a 2 segme e c u o o um wi a me ia im essio e y a s ia e egs s ou , e segme s sim e. A u s a e [ 8] Co ica ii ae: a i ii ae occasio a y ou u e a k o ea ee u ks. Co g. 208 En v p ou ec e o em e o Ma c . eco e om e e so a ea A sma , e o ga e o a , g ossy ack ee e . 2.0 mm (se e a eco s o I e ca gi . Io g. A e ae s aig , wi a segme e ack c u e y a i e a s co e , wi i e a s a cos a e [ 4] Ce y o i ae: Eu es i ae os e io y, a i e a cos a e oug ou . Occu s i g. 20 p d n ll r br p ( ei e ea i e a e ase o se ges. ou eque y i e A sma , o a , s o g y co e , g a ous ee e 2.0 .0 mm Auck a a ea, a a a e y wi e y is i u e . o g, a k ow a g ossy wi a e ages e is . ea s ig y i c i e , a owe a o o um a e ae [ ] Myce o agi ae: Myce o agi ae 0 segme e wi a 2 segme e ou c u , a e ice ig. 20 r ph ll h p d ll ( ou a sca e asymme ica o o um a e oi a , si ua e A sma , o a , s ig y e esse ee e .8 . mm o g, asa y e y a wi i e u c u es a ome wi s ack wi a e ae a egs ye owis i e, s o ack e i e ea i g emo a i es e i e wi i e a e a u esce ce a a e co ou e s o s o e y a. A e ae e a sio s. A u s occu i ea i e a o e o ga ic s e e , wi a segme e c u , e segme s sim e. i e . e omi a y om e o I., u some eco s ea e o swe om ma uka ( pt p r p r om e e so a ea. a o e s u s a so o us es. A u s we e o igi a y isco e e a W a ga ei ea s, a a e e o e om [ ] E omyc i ae: Myce aei ae e oises Is, Moko i au I., a se e a oca i ies i e ig. 204 M t bt rr n ( a icius e so a ea. A sma , o a , ai y, e is ee e . 2.0 mm Io g, wi s a se o u i g ai s. o o um si ua e asa y, wi a [60] A c eoc y ici ae im esse a e a i e o ei e si e e y a e o ga e, wi g. 2 0 Ar h r pt t p l Kas a is i c s ou e s. A a e i e s ecies, k ow as a mi o A sma , u i o m y o a , i e y u esce ee e 2. . mm es i em e a e egio s, ee i g o mou s i s o es a Io g, a k ow o ea y ack. A e ae wi a ce a s. eco e om e e so a ea. segme e c u os e a ocess a u y e a e a ica y. I o uce i o ew ea a om Sou Ame ica.
— — [6 ] Cii ae: Cii ae e a i e y assig e o is s ecies. eco e om o ig. 2 C z l nd ei e mai is a s. A sma , su cy i ica ee e .8 2.0 mm o g, a k ow , wi s o is es. A e io ma gi o ea wi 2 u [6 ] Co y ii ae: Co y ii ae ee a e ae wi a segme e , oose y o me c u ig. 2 6 h z n nt S a o o um oa y ou e a e io y, a o ima e y A sma , a ow y o a ee e . 2.0 mm o g, s ig y s aig a ase e y a wi co use ows o u c u es. e esse , wi u e su ace ea i g e se a coa se A u s a a ae a e o e ou oge e i cy i ica u c u es. A e ae wi a oose segme e c u asa u e s i a is s s e u gus (G n d r ppl n t , a e a segme u y isi e om a o e e y a mos y u a u s a so occu i e o es ca o y a ess su a a e egs s o . A u s occu i ac i es o ea eque y i i e . A ai y commo s ecies, wi e y is ea es o e ee e C th d lb t , a i ecayi g i u e . woo o oa ea e woo y a s. We a e see s ecime s om e o I. o y. [62] Me a yi ae: Me a yi ae ig. 2 2 l b n b l ou [66] U o i ae A me ium si e , e o ga e o a ee e 4. 4.4 mm o g, ig. 2 S rph t d r n t ascoe o a ge ow a iega e wi a k ow , wi s o e a A mo e a e y Ia ge, oa y o a ee e 8.0 .0 mm o g, o ge u esce ce. ea e u e o o um a e ae s ig y e esse , wi i egume ow , u a ia e i wi ou a is i c c u i egs s ou e a e o e s, s a e a co ou u e su ace wi a c es o sca es ea i g Io g, mi u e y se a e i ia s u s me aco ae a ge o mi g a a ia e a e . ea a ia y co cea e y a a . A u s may e ou a e i g o e u e si e o o o um eyes a mos ci cu a a e a i se io s e ecayi g ogs, w e e a ae o a y ee o u gi. ose a e ae s e e , wi ou a c u o o um wi eco e om o mai is a s, om ea sea e e o o a g es s o g y o uce a e io y e y a oa , mo a e a i a s. muc oa e a o o um. A u s may e ou c i gi g o e u e si e o ogs, u a e i icu o s o ecause o [6 ] Mo e i ae: Mo e i ae ei c y ic co ou a e . a ae i e i o e ogs. ig. 2 M rd ll nt r t W i e eco e om e Co oma e o M o e ea e so . A a ge i ai ee e mm o g, ack wi c a ac e is ic w i e ma ki gs o e y a. A e a s ig y se a e [6 ] C a co yi ae o o um si ua e asa y e y a ou e os e io y Ias ig. 2 8 Ch l dr v r t e e ac e a omi a segme o o ge i o a e mi a s i e. A u s A a ge, g ossy ee e 6 mm o g, a e o a k g ee o occu eque y o owe i g ma uka ( pt p r g ee is ow , o e wi me a ic e ec io s, a ow p r o a a (M tr d r r b t i u su , a e s o s o e y a, u co ou a e a ia e u e si e a k e y ac i e a goo um e s, a ake o ig ea i y g ee is o ow a e o emo a, i iae, a a si w e is u e . Wi e y is i u e . a ke e a e ae a e ow o sa su ace o ea a a e o ma gi s o eyes, o o a si es, a isc o [64] i i o i ae: e eco omi ae e y a wi sma a c es o coa se ye owis u esce ce. ig. 2 4 h p t n l br S a A e ae s e e , acki g a c u , wi i se io s e ose A me ium si e , e o ga e o a , ackis ow a e e e e e wee eyes ea i g a im u c a e a ea e y a ee e 6 8 mm o g. A e ae ame a e i ma e eyes su a a e o mos o e g egs s e e . Co i e o ki ey s a e , a y su ou i g ase o a e ae ig ai a o es s, mos y th f . eco e i e o o um wi si es a cua e a ase si ua e e y a wi o I. om e ce a o ca ic a eau a a ou We ou e a ices a e y co e i g wi gs, o s ia e, wi i g o , a i e Sou I. om Wes a a io a . e i i e o gi u i a i ges. A u s a e ee co ec e y ea i g ege a io i summe ey y e y ea i y. a ae [68] e e io i ae: ag ii ae a a e y a e e a o y, i ea woo . eco e om e ig. 2 Ch r d tr h l d W i e o I. a e o e Sou . A me ium si e , o us , s o g y co e ee e 6. 8.6 mm o g co ou a e e y a ia e, om u ico o ous [6 ] Co y ii ae: Co y ii ae a k ow o a wi a ke ma ki gs o o o um a ig. 2 r t d r nt r t W i e e y a. o i iae s o g y e a e a e o a e a y, wi A mo e a e y Ia ge, oa , a e co e ee e .8 8.0 a si su e mi a meso a me a i iae wi ai s o s ou , mm o g, wi mos o su ace ea i g s o is es. u , eg ike a ica s i es me a a si a g e u wa s a A e ae wi a segme e c u a e a ma gi s o ackwa s — a mo i ica io s o mo i g o oose sa . o o um wi 4 omi e ee e y a mos y su a a e , eco e om sa y coas s om e a o o S ewa wi ai y i egu a , o gi u i a y a a ge e e a io s I., o e u e seawee o o e eac w ack. a omi a s e i es u y mo a e. A u s occu o s a i g ea ee u ks a some imes u e oose a k, o e wi mou s. a ae ou i associa io wi a u s a e
— 8— [68] e e io i ae: ime ii ae qua a e e y a oa s ou e e , coa se y u c a e, ig. 220 A t z t lb t ascoe wi ou s ae. ou o o es g ow i summe . A u s A sma , a e co ou e a k i g ee e 2.8 .0 mm o g a e o e ee e s, a a ae o a y i e u e oose u e su ace co e e wi a e g eyis w i e sca es. a k o ea a c es. Wi e y is i u e om Auck a o ea a y co cea e y o o um eyes coa se y ac Sou a . e e o o um oa , wi o a g es oa a s ig y o uce a e io y e y a wi s iae o e egs oa y [ ] y oc oi ae: i i a i ae e a e o iggi g, a i egs ea i g o g is es, ig. 22 Ex l p p t n t ou a a a a io o wa ki g o sa . i es i oose sa o A me ium si e , e o ga e ee e 4. . mm o g, su eac es oug ou ew ea a . a a e si e wi e y a s ig y oa e i g os e io y, g ossy ack, wi i e u esce ce. ea a owe e i [68] e e io i ae: e e io i ae eyes, o mi g a mo e a e y a ow eck eyes omi e ig. 22 M p l n t ( me ma i a y a wi Ias segme oa y ia gu a a e A a ge, e o ga e o a , .mo e a e y co e a k i g ee e ae se a e i ema e, ec i a e i ma e o o um s ig y mm o g, ack o e is ack u e su ace a s e se, wi se e a im essio s e y a wi omi e c ose y u c a e, e i e s ices e wee u c u es s ou e s, coa se y u c a e, wi ou s iae. Occu s i a ia e su ace u o g ossy, wi o wi ou a a e o es e a eas o Gis o e, au o, a a aki, We i g o , mic oscu u e. ea wi e a e ma gi e ow eac a e so o Sou a , mai y a ig e a i u es. eye o mi g a ca us a e ae wi a i is i c segme e c u o o um a s e se, e o a g es [ 2] Sa i gi ae: Sa i gi ae o uce , wi ou s iae egs s e e . A u s usua y ee ig. 226 S lp n b l n t ascoe o ea ege a e e is, u i y co i io s ee o A sma , e o ga e o a ee e 2.0 2.2 mm o g, wi a i i g a issue. Wi e y is i u e , a icu a y i coas a is i c wais e wee o o um a e y a, ye owis oca i ies u e c ee i g coas a a s suc as o ue ue. ow wi ack s o s o e y a, e s o s a ia e i Auck a a o some o s o e is a s commo u e s a e, a some imes use oge e . A e ae wi a s o es i c a e s o e i c o ca oes. ackis 4 segme e c u o o um wi a ge, ee u c u es e y a wi ou is i c s iae, u wi s ig y [6 ] os omi ae i egu a ows o coa se u c u es. A u s eque y ou ig. 222 r r h tt ascoe y ea i g sc u y o es g ow i summe . Wi e y A me ium si e , e o ga e ee e 6.0 8.0 mm o g, su is i u e . a a e , e esse , usua y e is ow . ea wi os e io y o uce o es e i eyes o o um e o [ 2] Sa i gi ae: I o e i ae ga e e y a ai y u c a e s ia e. A u s a a ae may ig. 22 r pn d f ld Wa e ouse e ou i ogs o imu a i au, i am c e ices e A mo e a e y a ge, e o ga e, s o g y e esse ee e wee woo ocks i e mois , ee e s age o ecay. .0 0 mm o g, g a ous, a e ow wi a ke e y a. Wi e y is i u e . ea e o ga e e i sma eyes a e ae i i o m, wi sca e e a ge o o um s o g y a owe os e io y, [ 0] Oe eme i ae: Oe eme i ae u ca e a ica y e y a s o e e , s ig y co e , wi ig. 22 h l ph l n t ( a icius a e oa y ou e , ea i g a eas 4 a omi a seg A a ge, e o ga e o a a ee e 8 20 mm o g, s ig y me s e ose . ou wi a ae, a so e y e esse , e esse , wi i egume so . ea o uce i o o u e oose a k o ea u ks a a c es, ee i g o eyes a e ae o g, a e ye owis ow , wi ou a c u , ecaye cam ia issues. Wi e y is i u e , a eco e e segme s su cy i ica , wi sca e u y e ose om e C a am Is. o o um wi a a k me ia s i e e y a wi a k a e a o gi u i a s i es, coa se y a c ose y u c a e, wi 2 [ ] A ici ae: A ici ae i is i c o gi u i a cos ae. A u s a e ou o owe s o ig. 228 Anth h p r Ki g a i e s u s, w e e ey o a y ee o o e a A me ium si e a ee e . 4.0 mm o g, ack wi ec a . Occasio a y ey occu i o e woo o e o es o a ge ye ow e y a s o s a os e io o o um oo . a ae i e i we o e woo . Wi e y is i u e , wi u esce ce s a se. ea su a a e e i eyes, e eco s om S ewa I. a e C a ams. a u y co s ic e i o a a ow eck eyes sma a e ae wi ou a c u ma i a y a s wi asa 2 [ ] y oc oi ae: i i a i ae segme s sma a cy i ica o o um a owe ig. 224 h n l r a es os e io y e y a wi o ou ce s ou e s, a mos wice A me ium si e , e o ga e, su a a e si e ee e .0 o o a wi me aco ae se a a e y mo e a o e 6.0 mm o g, s ig y e esse , co e e wi i e u co a wi . eco e i Auck a om a a ock, esce ce. ea a owe e i eyes, o mi g a oa ea e u coa se u i gs, a a s eam e i e us . eck eyes omi e a e ae se a e Ias segme o A so k ow om e so , a o a y wi e y is i u e . ma i a y a oa y ia gu a o o um sma , su
- - [ ] A ici ae: ag ioi i ae mm o g, s ig y e esse , wi o y so a agi e, ig. 22 r d br n ascoe u esce , a e ye ow wi a k ow ma ki gs o e y a. A me ium si e a ee e 4.0 . mm o g, o a ge wi ea s ig y a s e se, e e e eyes C s a e , a y e is ea , o ea a o o um ow wi e y a su ou i g a e a ase a e ae i i o m ma i a y mo e ye ow a ow egs a a e ae ye ow o a s wi as segme oa y e a e o o um o a ge a e a c u some imes a ke . Associa e wi s o g y a s e se, wi 2 sma asa im essio s e y a coas a sa u es, a may e ou e ea eac wi co use u c a io , Iacki g s iae i ia s u s sma , e is. eco e om o mai is a s, S ewa I., a wi mi u e u esce ce. eque y ou o o iage a e C a ams. owe s i summe . Wi e y is i u e o o mai is a s.
[ ] A ici ae: emo i ae [ 6] Ce am yci ae: io i ae ig. 2 0 C t r p ( ou g. 2 4 r n pl r t l r W e A mo e a e y sma a ee e 2. . mm o g ea , e ea ies ew ea a ee e, 40 0 mm o g, oug o o um, a ase o e y a e is emai e o e y a ike mos o e woo o e s a ia e i si e e e i g o o a ge, wi a ow a s e se s o me ia y a e e u i io o e a a, a k ow wi a e icu a e ages ye ow, g ossy u esce ce mo e a e y e se a ye owis a e o e y a. o y e o ga e, somew a ye ow. ea a ow y co s ic e e i eyes i o a e esse a e ae a mos as o g as o y o o um a ow eck eyes a ge, o u i g a e ae i c assa e wi a e a ma gi ea i g s i es e a su ace co e e a ica y u wi ou a is i c c u asa 2 segme s o wi e se u esce ce egs o g a s e e . a ae ma i a y a wi sma o ec io s o o um co s ic e a ack mai y ea co i e s, es ecia y a ia a i e a ea ase e y a wi ou is i c s ou e s, ess a wice imu, u a e e o e a so om wa e ey equi e woo o o a wi . Co ec e u e h r t n x, A t l wi a ig mois u e co e . A u s o o ee , u i isi g a b n , a G hn tίf lί . O igi a y esc i e om ese es ai ow y e a ae. u u g u s we e o ce a a au, a wi e y eco e i e o I. sig i ica oo esou ce o Mao i. eco e om e Ke ma ec Is o S ewa I., o e a ig . [ ] A ici ae: Mac a ii ae ig. 2 M r tr x l ascoe [ 6] Ce am yci ae: Ce am yci ae A me ium si e a ee e 4.0 4.2 mm o g, a k ow o ig. 2 r n s . ea y ack wi asa a e a segme s a egs A sma o g o 6.0 .0 mm o g, e is ow wi e is ow o o um a e y a coa se y u c a e. ye owis s o s o e y a, occu i g o e Ca e u y ea a u y co s ic e e i eyes i o a a ow eck ai s. e e a e se e a e emic r n s ecies i ew eyes Ia ge, o u i g a e ae wi a oose y o me , ea a , a sma a s iki g y co ou u , u esce o i is i c , segme e c u ma i a y a s wi asa a mos g a ous (e.g., ue wi o a ge s o s, o a ge wi segme s ea i g ia gu a o ec io s o o um o a , ye ow s o s , u ecause o ei sma si e ey a e wi ase a a e oa e y a wi o ou ce eque y o e ooke . A u s ca e ou ee i g i s ou e s, a mos wice o o a wi me aco ae a mos a i e owe s. eco e om o mai is a s. ouc i g. ea e om o iage i mi summe , a ou o se ges. eco e om ea sea e e a ou e o I. [ 6] Ce am yci ae: Ce am yci ae ig. 2 6 O n h rt ( a icius [ 4] A e i ae A a ge, su cy i ica o g o ee e 2 mm o g, ig. 2 2 ph l n t d ou ma a k ow wi scu e um a eyes o a ge. A e ae A sma , e o ga e o a , s ig y e esse ee e . 2.0 s e e , as o g as o y o ea y so o o um wi mm o g, ack wi egs a a e ae ye owis , u c a e, i egu a a s e se g oo es e y a wi ou s iae, co e e c o e wi g eyis ecum e ai s. ea s o g y wi coa se ow is u esce ce. A u s co ec e y a owe e i eyes i o a is i c eck eyes omi e , ea i g o es g ow u i g summe , o occasio a y a coa se y ace e a e ae s ig y oa e i g a ica y, ig . a ae a ack ma y woo y a s, o i g i o i e wi e mi a segme e a ge a o a o o um sma , woo o ci us ees ( e ce e commo ame emo ee a o e y a wi e y a oa , wi a e essio e i o e , g a e, a a ious a i e a s, es ecia y s ou e s, u c a e u wi ou s iae i i iae wi a gio a. eco e mos y om e o e a o e s a a ica ocesses. A u s occu o o iage o M r t o I. a a ou e so i e Sou I. n l r ( ee Ki gs uka a o e s u s, a o e s ecime was ou i a o i s es . isco e e a [ 6] Ce am yci ae: amii ae o co e, Auck a , a su seque y eco e om ig. 2 l t l t p nn ( eu i g y ie a om a y A ice I., e a C icke s g ou . A mo e a e y a ge, a ow y o a , co e o g o ee e 8.0 4 mm o g, wi o o a cy i ica , su g a ous, [ ] Sc a ii ae a e ow o a k ow wi a c es o a e , ye owis ig. 2 th t l t t ( ou u esce ce o si es o o o um a 4 a ow y e o ga e A mo e a e y sma , e o ga e o a ea i g ee e 2. .0 s o s o e y a. ea o ie e ow wa s eyes s o g y
80 ema gi a e a e ae o ge a o y, i se e i [ ] C ysome i ae: C y oce a i ae ema gi a io o eyes o o um cy i ica e y a wi ig. 242 Arn br n S a i co s icuous s ou e s, oa es i asa a egs s ou . A mo e a e y sma , su cy i ica ee e 2. . mm o g, eco e mai y om Auck a , Co oma e , a o us ow wi 6 a ica a e a segme s a ke , a a . o e a ke ea , egs, a o y ma gi s e se y a coa se y u c a e. Eyes e e y co e o o um ema [ ] C ysome i ae: uc i ae gi a e a wi ou ee ygi ium usua y a y e ose . ig. 2 8 r h d v ll ( a icius Co ec e y ea i g ma uka ( pt p r p r A sma , o a e ea ee e . . mm o g, u esce , i owe . eco e om a ou Auck a , a i e s ig y g ossy. ea a mos e i e y co cea e y o e so a ea. o um a e ae wi Ias 4 o segme s e a ge o o um semici cu a a e io y, si ua e asa y a wi [ ] C ysome i ae: Eumo i ae u esce ce o mi g si ua e a e s i me ia a o isc ig. 24 E l p br nn ( a icius e y a oa a s o e e , e osi g ygi ium, wi A me ium si e , e o ga e o a ee e 4.8 .0 mm o g, is i c s iae me a i ia wi a i co s icuous os e io ye owis ow u a ia e, a gi g om a k ow o s i e. I o uce om e U i e Ki g om o co o o ow is oc eous, usua y wi a ackis s i e ow e oom (C t p r . su u e a o e o ei e si e o e y a. U e su ace s o g y u c a e a e ae i i o m. Commo o ma uka [ ] C ysome i ae: C ysome i ae ( pt p r p r , es ecia y w e owe i g, ig. 2 Chr l n h p r o s e a o e o iage o o e a i e ees a s u s. o e A me ium si e , oa y o a ea ee e .0 .0 mm o g, ee e a ae i a i soi , es ecia y u e ma uka. Wi e y me a ic g ee o ackis me a ic g ee . ea ee y is i u e . i se e i o o o a a e ae s o , oa y se a a e asa y o o um a s e se, s ig y ema gi a e a ica y, [ 8] emo yc i ae: i o y c i ae si ua e asa y, a mos equa i wi o e y a e y a oa , ig. 244 h n rh n h r f l ( ou ou e os e io y, wi i e, e se u c a io a a ge, A sma wee i 2.0 2.8 mm o g, a ow y o a , us ow s a se u c u es i ou e co use ows egs s ou , wi o a k ow wi mo e a e y e se, ecum e , ye ow a i iae simi a y e e o e . I o uce i o ew ea a is g ey u esce ce. ea a u y co s ic e i o o o io ogica co o o S o s wo ( p r ,a eyes i o a a ow, a ica y a e e os um a e ae ow es a is e . i se e i a ica a o os um, s aig , wi a oose segme e c u o o a as o g as wi e o o ge i [ ] C ysome i ae: Ga e uci ae (A ici i ma e, as wi e as o g o s ig y wi e i ema e e y a ig. 240 A l h r ph l Se ma & og is i c y s ia e i wi gs we e e o e . A u s a e A me ium si e , e o ga e o a ea ee e 4. .0 mm associa e wi o oca acae, a a ae a e o e o g, ack wi ase o a e ae ye owis e , a wi ee e s i ma e i o esce ces o o oca s. eco e o gi u i a U s a e w i is ye ow s o s o e y a u e om e a o o S ewa I. su ace e se y u c a e, e u c u es co use . A e ae a o ima e asa y o o um su qua a e, [ ] A i i ae: A i i ae is i c y a owe a e y a a e a ma gi o o o um ig. 24 Sh rp v n t ( ou a me ia ma gi s o e y a su a a e e y a wi is i c A mo e a e y sma , e o ga e o a u gus wee i . .4 s ou e s egs o us me a emo a swo e . A Sou mm o g, u esce a sca e i egume ack wi Ame ica s ecies i o uce i o ew ea a om Aus i ia c eam a o a ge ma ki gs. ea wi os um a ia o io ogica co o o a iga o wee (Alt rn nth r o ge a wi e, e a e a ica y a e ae s aig , ph ll x r d , a ow we es a is e i o a . wi a segme e c u , e asa segme symme ica a segme 8 e o ga e a s ig y a owe a ica y [ ] C ysome i ae: Ga e uci ae (Ga e uci i o o um s ig y o ge o s ig y s o e a wi e, wi ig. 24 Ad x v l r ( ou a s e se ca i a s ig y cu e a a e a ca i a s o A me ium si e ee e .0 6.0 mm o g, wi ea , o ygi ium ugose i ma e a u c o as e a e i ema e. o um, a e y a u i o m y ye owis us ow , o ea ea e om a ious a i e a i o uce s u s. Wi e y ow , o o um o a ge ye ow, a e y a o i e o ye ow is i u e o o mai is a s, om o oe u S a e is ow wi a k ow e ges a e ages usua y a k o es o I e ca gi . ow . U e o y mo e a e y co e , g a ous o ea y so, e se y u c a e, e u c u es i e ea a [ ] A i i ae: C o agi ae o o um is i c y a owe a e y a. Co ec e om ig. 246 n r pt p ll d ou o e a c es (e.g., d rp , i o esce ces o A sma , o a , u esce u gus wee i .2 2. mm o g, a gio a ( r h l tt r p nd , a ea e om s u s. a iega e e is ow a ye ow, o e wi ack o Wi e y is i u e ac oss e o I. a o e Sou I. a k ow ma ki gs, a e y e i e y ack. ea wi a s o , oa os um a e ae s aig , wi a seg
— 8 — e e c u , e asa segme asymme ica o o um ow o ow o a mos ack wi s a se, mo e a e y wi e a Io g, wi a s e se asa ca i a e mi a i g o g, ye owis g ey u esce ce. ea a o o um wi a u y a a e a ma gi , a a e a ca i a a se e y a mes e mic oscu u e os um e o ga e, a ow a e e o ga e, wi s iae co ai i g a ge u c u es. A u s ae s ig y ge icu a e, i se e ea mi e o os um, a e ee co ec e om ea i e a ea e om ea wi sca e a mos as o g as emai e o a e a a o eysuck e. is i u e i e sou e a o e o I. c u segme e o o um a owe a e y a, s ig y a o eas e Sou I., i c u i g some o s o e is a s e o ga e a co ac e a e io y e y a o a i ou i e, (S e e s I., U i e ., a C e wo e Is . wi coa se y u c a e s iae a co e i e s ices. A u s occu o o iage o o u ukawa (M tr d r x l , [80] e i ae: e i ae a a ae i e su co ica y i a c e s. eco e om ig. 24 C r t ph tr d n ( a icius o a o Auck a a e Co oma e a ea, i c u i g A a ge, i egu a y s a e wee i 0 mm o g, a e e A e me , Me cu y Is, i e a ie I., O e a I., e ow , mos y co e e wi w i is a i k sca es. ea a C icke s Is, Co e mi e I., a e Moko i au Is. a o o um a owe a e y a a a e i g a e io y ea o co s ic e e i eyes, wi os um mo e a e y [82] Cu cu io i ae: ac yce i ae e o ga e, oa eyes omi e a e ae s aig , wi ig. 2 An t t rb tt (S i e ou a c u o o um a e oi a e y a oa , u e cu a e, A a ge wee i u o 24 mm o g, ow o e is ow a owe su a ica y a sa segme s o g y o e . wi w i is o sa s o s, we ma ke o mi e a A u s a e eque y ou i ea woo o ma y ki s. os e io e y a e a su ace w i is . E y a wi a ge, eco e om e o I. a o e Sou I. co e s a e os e io o u e a ces. A u s may e ou i e a c es o a i e ees (e.g., gaio , w e e [8 ] e i ae: Ag ycy e i ae a ae a e mos ike y woo o e s. Su i i g o y o a ig. 248 Ar l ll t n (S a ee o s o e is a s suc as e ee Ki gs, oo A mo e a e y sma , e o ga e, su a a e wee i .0 .2 K ig s, a Mu iw e au i e e a C icke s g ou . mm o g, u ow , u esce . ea a u y co s ic e e i eyes, wi a u os e io a g es os um s o , [82] Cu cu io i ae: ac yce i ae oa , s o e i ema es a i ma es a e ae s aig , ig. 2 2 M nd l t r ll ( ou wi a is i c 2 segme e c u o o um a o ima e y A me ium si e , a ow y o a wee i 4.0 .0 mm o g, su qua a e e y a s ia e u c a e a si wi 2 seg ma ow is g ey, usua y a iega e wi i co s icuous, me s o g y o e . A u s a a ae may e ou sma , i egu a y s a e a e s o s. o o um a e ea a k o ea o yi g a c es o i e i ge . a icu a y e y a wi sca e e e o ga e sca es os um A a e y wi e y is i u e . oa , s o a e ae s o g y e owe o o um as oa as e y a, wi mo e o ess is i c a s e se [8 ] e i ae: e i ae w i k es e y a wi o gi u i a s a ow i ges. Occu s o ig. 24 rh n h b rb rn ( a icius sa y eac es a coas a ege a io , oug occasio A e y a ge, e eme y a ow y e o ga e wee i 8 a y ou i ga e s i c us e s o a . eco e i e mm Io g (ma e ema e 8 4 mm , u c a e, wi a ge Auck a a ea. u c u es o e y a mi u e ye owis u esce ce o ea , o o um, a e y a a a i io a ow a o ge [82] Cu cu io i ae: Cu cu io i ae u esce ce o os um, a e ae, a a si. o y ma ig. 2 Andr ll p r ( ou ow a iega e wi e is ow s o s, i some A sma , o a wee i . 2.4 mm o g, ma ow , wi s ecime s a mos u i o m y a k ow os um e eme y ow a a e ow o ec i g sca es. os um a ow, e o ga e, s o e i ema e a i ma e a e ae s aig , o g a e ae is i c y ge icu a e o o um s ig y i se e a ica y i ma es, me ia y i ema es o o um a owe a e y a e y a wi sca es o mi g i egu a wi a me ia su cus e y a co s ic e su a ica y i ows egs s ou , wi o ec i g sca es. A u s a e ee ma es, e e y a e i g i ema es a si o g. O i osi io co ec e y ea i g A t l s . a usk a so ou o si es a e e a e y ema es om Oc o e o Ma c y o u ukawa a i yo y es i e se o es . eco e c ewi g a ca i y i e a k o yi g o su esse ees om o a , Auck a , a e ay o e y. (kau i, igeo woo , ewa ewa, awa, ka aka o ogs. ew y a c e a ae o e a ia y i o e woo , a [82] Cu cu io i ae: Cu cu io i ae e a ge e u e as ey g ow. Am osia u gi a yeas ig. 2 4 M rt n z l nd Kusc e g owi g i e u e s mos ike y se e as a a oo . A mi u e, a ow y e o ga e wee i 0. 0.8 mm o g, a e eco e om e o I. a o e Sou I., mos y a ye owis o e is ow wi a e ages usua y a owe a i u es. s a e a e . os um mo e a e y e o ga e, s a se y u c a e eyes com e e y e uce o o um o ge a wi e, [8 ] e i ae: A io i ae a ia y a s a se y u c a e e y a a e e , wi i e ig. 2 0 b tr d r ( ou u c a io o mi g o so esce s iae. As a as is k ow , A sma , y i o m wee i .8 2. mm o g, g ossy e is is i s ecies is e sma es wee i i e wo .
— 82— Co sis e y ou a ou e i e oo e s o a a i e [82] Cu cu io i ae: Sco y i ae s ecies o My aceae, i occu s i e o 0 cm o soi ig. 2 8 l t t r ( ayku u i g we e co i io s a a g ea e e , 0 60 cm, A me ium si e , a ow y e o ga e wee i . 6.0 mm u i g y e io s. eco e om e o e o I. o g, a k ow o ack, a e y e is ow a e ae a a si usua y e is ow . o o um e o ga e a [82] Cu cu io i ae: Cu cu io i ae e se y u c a e, e u c u es ou e y a u c u es ig. 2 St ph n rh n h l n S a ou , a o mi g s iae, wi s o i co s icuous A sma , is i c i e wee i .2 4.8 mm o g, c a ac e ise u esce ce. ou mai y i i us ia im e a u e y i s o g, ee y co s ic e ea , co ica o o a , a k o n r d t ogs. eco e om o mai u e cu a e e y a, a e ao i a i y o g i egs. ese is a s, a gi g om Wai oua S a e o es o O ago. a e s o g y cu e i iae a emo a wi a a ge, ec a gu a oo , e a i g e wee i o c as u c es o [82] Cu cu io i ae: Sco y i ae owe s ame s a ake o e owa s e mou . ig. 2 hl n pr o ki s Commo i o a a e o e Sou I. A sma , mo e a e y s o wee i 2.8 .0 mm o g, su cy i ica wi o e o y s ig y a e i g a e io y, [82] Cu cu io i ae: Cosso i ae su g a ous wi i co s icuous s o a e ec se ae, ig. 2 6 M r t l r t (S a mo e a e y g ossy. ea a ge y co cea e y o o um A sma , a ow y e o ga e wee i 2. .0 mm o g, g ossy om a o e os um s o a oa o o um a owi g ye owis e o a k e is ow su g a ous, wi a e io y, as oa as e y a a ase e y a mos y su i co s icuous s o a e ec se ae. os um o g, oa , a a e , wi ee , coa se y u c a e s iae. A a e i e s ig y e a i g a ica y a e ae s o g y ge icu a e s ecies eco e om Auck a o C pr t r l . o o um a mos as oa as e y a, ou e a e a y e y a mos y su a a e , wi a gu a asa a g es a [82] Cu cu io i ae: a y o i ae wi is i c u c a e s iae. ea e om a ious ees, a ig. 260 l t p p l W i e ea e om a ae i C pr r rp . i s e o e A me ium si e , su cy i ica , a ow y e o ga e wee i om Auck a . 6 mm o g, su a a e si e , g ossy e is ow o a k ow . os um e y s o a e ae weak y ge i [82] Cu cu io i ae: Cosso i ae cu a e, wi a oa c u o o um s ig y e o ga e, ig. 2 n n p n p Wo as o ec a gu a , s a ow y ema gi a e me io a e a y e y a A somew a a e e a e o ga e wee i .0 4.8 mm su a a e , i e y s ia e a si o g a s e e , wi asa o g, wi a s o , oa os um, e o ga e o o a , e y a segme s o g y e o ga e. A u s a e u y wi ge , a wi we ma ke s iae a u c a io , a ea y egs wi may e co ec e usi g Ma aise a s. ey o e i o a wi e s o g ee a e a e o e i iae. A u s a a ae co a ge o a i e a e o ic ees, maki g u e s a ou 2 s uc ga e ies u e oose a k o A th a Ar r mm i iame e a ig a g es o e g ai . ese ecome s ecies. es ic e i is i u io y e a ai a i i y o os i ec e wi am osia u gi, o w ic e a ae ee . ees, a e ce eco e mos y om o e oca i ies. Occu s wi e y o o mai is a s a o e C a am Is.
8 A E I : E a ge e ee e s ecies A E I 2: Ma o e omo ogica co ec io s [modified after Molloy et al. 1994] i ew ea a A number of species are as yet undescribed, and are AMNZ Auckland Institute & Museum referred to by informal names or by `sp.' Synonyms and Private Bag 92-018, Auckland superseded names are in parentheses. Attn: Mr John W. Early, Curator of Entomology Tel: +64 9 309 0443 Fax: +64 9 379 9956 • Seven species are listed in Category A (highest priority Internet: [email protected] threatened species): Geodorcus (=Dorcus) auriculatus, G. ithaginis, G. 'Moe- CMNZ Canterbury Museum hau', Mecodema costellum costellum, M. laeviceps, Pro- Rolleston Ave, Christchurch dontria bicolorata, P. lewisi Attn: Simon Pollard, Curator of Invertebrate • Ten species are listed in Category B (second priority Zoology Tel:threatened +64 species): 3 366 8379 Fax: +64 3 366 8379 Internet: [email protected] Amychus granulatus, Anagotus stephenensis, A. turbotti, Hadramphus spinipennis, Heterexis seticostatus, Lypero- FRNZ Forest Research Institute bius huttoni, Megadromus 1 sp., Oregus inaequalis, Pro- Private Bag 3020, Rotorua dontria grandis, Zecillenus (=Cillenum) tillyardi Attn: Roger Crabtree, Curator of Entomology Tel:• Six species +64 are listed in category 7 C (third347 priority 5899 Fax: +64 7 347 5333 threatened species): Internet: [email protected] Amychus candezei, Anagotus fairburni, Hadramphus stilbocarpae, Mecodema chiltoni, Oclandius laeviusculus, LUNZ Entomology Research Museum Prodontria modesta Dept of Entomology & Animal Ecology, P.O. Box 84, Lincoln University, Canterbury • n p r l t d n C t r Χ ( p h h h v Attn: Mr John W. M. Marris, Curator Tel:not been sighted+64 for a number of3 years but325 which may 2811 Fax: +64 3 325 3844 still exist): Internet: marris @tui.lincoln.ac.nz Hadramphus tuberculatus, Mecodema costellum `spelles', M. punctellum, Megacolabus sculpturatus, Megadromus MONZ Museum of New Zealand Te Papa Tongarewa antarcticus 2 subspp., Pericoptus nitidulus, Prodontria Natural Environment Dept, P.O. Box 467, Wellington `Five Rivers', Stethaspis convexa, Thotmus halli Attn: Mr Ricardo L. Palma, Curator, Terrestrial Arthropods Tel:• Al t ht +64 p r n d4 t 381 C t r Ι 7361 Fax: +64 4 381 7310 (species about which little information exists, but Internet: [email protected] which are considered threatened): Amychus 1 sp., Blosyropus spinosus, Brullea antarctica, NZAC New Zealand Arthropod Collection carabid `Kamo', Euconnus microcilipes, E. paracilipes, Landcare Research, Mount Albert Research Centre, Holcaspis falcis, Lyperobius carinatus, Maorinus hunu- Private Bag 92-170, Auckland aeformis + 1 sp., Maoripamborus fairburni + 1 sp., Meco- Attn: Dr Trevor K. Crosby, Curator Tel:d ll n +64 M n t l M 9 tr x M 849 br tt n Μ 3660 Fax: +64 9 849 7093 t ll l M t ll b M rv d n Μ Internet: [email protected] dunense, M. dux, M. howitti, M. integratum, M. litoreum, M n x M r M n t d M p v d Μ pl t Μ OMNZ Otago Museum pr x M p l h ll Μ n n Μ r x Μ P.O. Box 6202, Dunedin strictum, Megacolabus bifurcatus, M. obesus, Megadro- Attn: Mr Anthony C. Harris, Curator of Zoology Tel: p t (=b l +64 M pr 3 M 477 f lt n Μ 2372 Fax: +64 3 477 5993 haplopus, M. `Omarama', M. v r n + 5 spp., Metablax 1 ρ Mimopeus parallelus, Nesoptychias simpliceps, Note. New Zealand beetles may be found in institutional Neuraphaconnus toronouii, Nothaldonus peacei, and private collections around the world. For instance, the Oclandius cinnaraeus, Paralissotes mangonuiensis, Broun Collection with its many primary type specimens is Paratorchus alifer, P. flexuosus, Pericoptus frontalis, held at the Natural History Museum, London, and the Pheloneis gratiosus, Prodontria patricki, P. regalis, P. Field Museum, Chicago, has extensive holdings of Staph- setosa, P. `Twizel', Sciacharis yakasensis, Stephano- ylinidae and Leiodidae from New Zealand. Despite name rhynchus insolitus, Syrphetodes 1 sp., Tangarona pen- changes affecting some institutions, the list of repositories sus, Tychanopais tuberosus, Zeopoecilus 5 or 6 spp. published by Watt (1979c) is still a useful introduction.
—184— A E I Key o i es o Ca a i ae i ew ea a [provided by J.F. Lawrence]
1 Clypeus broader than distance between antennal sock- - Elytra with an inner longitudinal ridge, its end visible ets; eyes large and protruding; protibia with 2 spurs beneath edge of elytron, level with apical end of terminal; lateral pronotal carinae absent or incomplete epipleuron; head with frontal furrows not extending posteriorly ... Cicindelini behind eyes ... 10 —Clypeus narrower than distance between antennal sockets; without other characters in combination ... 2 10(9) Maxillary palp with penultimate segment setose; male protarsi with 2 segments dilated and dentate on 2(1) Metepimeron not visible between posterior edge of inner face ... Zolini metepisternum and anterior edge of 1st ventrite ... 3 —Maxillary palp with penultimate segment glabrous; —Metepimeron visible as a lobe between metepisternum male protarsi unmodified, or only slightly so and 1st ventrite ... 5 Psydrini
3(2) Procoxal cavities closed behind; apical segment of 11(7) Elytra with apex rounded as a single curve, or maxillary palp slender and fusiform; length less than acutely angled at apex; Abdominal apex usually 15 mm ... Migadopini concealed from above; cloration rarely yellow and —Procoxal cavities open behind; apical segment of black ... 12 maxillary palp more or less expanded and truncate —Elytra with apex transversely or obliquely truncate, apically; length more than 15 mm ... 4 exposing abdominal apex from above; coloration often yellow and black; body often flattened, or with 4(3) Mandibles multidentate, smooth on upper surfaces; prothorax elongate and narrow ... 15 clypeus without setiferous punctures ...Cychrini —Mandibles not toothed, rugose on upper surfaces; 12(11) Head with a single pair of supraorbital setiferous clypeus with a setiferous puncture at each anterior punctures ... Harpalini angle ... Carabini —Head with 2 pairs of supraorbital setiferous punctures ... 13 5(2) Body pedunculate, with a distinct, narrow waist or peduncle between prothorax and elytra; scutellum 13(12) Clypeus emarginate, with a pale membrane bet- entirely contained within peduncle, not forming a ween clypeus and labrum (usually within emargination); wedge between elytral bases ... 6 mandibles characteristically bifurcate, with a large —Body not pedunculate, or if slightly so then scutellum mesal tooth; labrum often deeply emarginate extending behind peduncle and forming a wedge Licinini between elytral bases ... 7 —Clypeus not emarginate, without an obvious pale membrane between it and labrum ... 14 6(5) Mesocoxal cavities open laterally, partly closed by mesepimeron ... Clivinini 14(13) Elytra with an inner longitudinal carina, its apical —Mesocoxal cavities closed laterally by meeting of end visible below lateral edge towards apex, causing sterna ... Broscini end of epipleuron to appear twisted; carina sometimes concealed by lateral edge of abdomen, which fits into 7(5) Mandibles with a setiferous puncture in concavity a groove between it and epipleural apex; mentum (scrobe) along outer edge ... 8 usually bifid at apex ... Pterostichini —Mandibles without a setiferous puncture in concavity — Elytra without an inner carina or, if present, not ex- (scrobe) along outer edge ... 11 tending to lateral edge; mentum usually simple or with a single median apical tooth ... Agonini 8(7) Maxillary palp with terminal segment very small Bembidiini 15(11) Mentum supported on a projecting submentum; —Maxillary palp with terminal segment usually about as head not sharply constricted behind eyes; mandibles long as penultimate segment ... 9 with hollow outer face; claws often pectinate ...Lebiini 9(8) Elytra without an inner longitudinal ridge beneath —Without a projecting submentum; head sharply apical edge; head with frontal furrows extending constricted behind eyes; mandibles without hollow beyond posterior edge of eyes ... Trechini outer face; claws simple ... Pentagonicini
—185- A E I 4: is i u io o co ec io eco s o ee e ami ies i ew ea a (Nth, North I. Sth, South I.; Stw, Stewart I.; Ke, Kermadec Is; 3K, Three Kings Is; Ch, Chatham Is; Sn, The Snares Is; An, Antipodes Is; Bo, Bounty Island; Au, Auckland Is; Ca, Campbell I. .
Family Sth S w Ke Κ C S An Bo Au Ca
Aderidae • Agyrtidae • Anobiidae • • • • • • Anthicidae • Anthribidae • • Archeocrypticidae • • Belidae • Bostrichidae • Bothrideridae • Brentidae • Buprestidae • Byrrhidae • Cantharidae • Carabidae • Cavognathidae • • Cerambycidae • Cerylonidae • • Chaetosomatidae • Chalcodryidae • Chelonariidae Chrysomelidae • • Ciidae • Clambidae • Cleridae • Cocci e i ae • Colydiidae • Corticariidae • Corylophidae • Cryptophagidae • Cucujidae • Curculionidae • • Dermestidae • Derodontidae • Dryopidae • Dytiscidae • Elateridae • Elmidae • Endomychidae • E o y i ae •
— 86— l th Sth St K Κ Ch Sn An A C E n t d E n d • . G r n d . t r r d t r d dr n d dr ph l d b n d phl d . • n r d • • d d n h d n d • • d M l ndr d M l r d M n t d M rd ll d • M t ph d n h d t d l d d ndr d O d r d h l r d hl t h d h d • . r t d t l d t l d t l d r hr d h p ph r d h d d . S lp n d S r b d S rt d S r pt d S d n d S lv n d St ph l n d n br n d r d r t d Ul d d — 8 — A O OMIC I E O y co eo e a a a a e i e e , si ce o e o ga isms me io e i e e , e.g., os a a, a e a ge y e i e om o e u is e sou ces. ames a e is e i a y og a ica ie a c y, as o ows. CA I A S: a a a o e ami y e e . o : e 82 ami ies a occu i ew ea a . o ma : e o ic ami ies, ami y sy o yms, su ami ies, i es. I t l : ge e a, s ecies. age um e s o w ic a a o is keye ou a e su i e k . o y e e o es e age o w ic a ami y is esc i e , a I a ic e age o w ic a a o is i us a e .
A aei ae 24 Anth 6 : h p r 6 , , A nth l d 06: bt t 2 A e i ae 40 A r d t n , , Anthr n r 40: tr l 4 Ac o ic i ae 2 Anthr n 40: v rb 40 A t z t 60: lb t 6 , , A i i ae 6, 68, k, 06k, 2 , 64, 86 A e ii i 6 A i i ae 68, 6 k, 64 A E AGA , , 8, 20, k, 0 k, , 4, 2 , A o ii ae 0, k, 22, 26 Apht r 60 A e i ae 64, 0 k, 0 k, 60, 86 Ap n 24 Ad x 2 : v l r 6 , 6 , 8 A io i ae 6, 6 , 0k, , 06k, 6 Adr t l r 26, Ar l ll t n 6 , 2 , 64, 82 Aegia i i ae 6 Ar h l n 8: t p nn Aesa i ae 2 , A c eoc y ici ae , 6, 02k, 0 k, , 86 Aga y i ae , 4 k, 2k, 0 Ar h r pt t p l 6, , A p th 0 : f v ll 4 , , Ar h l n 60 A l h r ph l 6 , 62, 8 A C OS EMA A , , 22 A th n tr d n 24, 2 Arh p l tr t 6 Ag ycy e i ae 6 k, , 8, 06k, 64 Ar d n d f r A n th t v n 2 , 6, 0 Arn 2 : br n 6 , 6 , 8 Ago i i 8 Arp d r n 2 Ag y i ae , 44 Arth p Agy i ae 2, 2 , 0 k, k, 2, 86 Arthr nth 4 Ah v r 48: dv n 48 Arthr l p 4 Al p t 2 Art t n 6 : r h n Al h r 6, 2 : h nd 2 , , 6 A t d r 2: b l t 2, , p nt r 2 A eoc a i ae 6, 2 , 28k, A l 8: f lv th r x 8, 4 , 4 f d 24 A ecu i ae 60, 6 , 26 A ph r t 26 All h r 2 A oma ii ae 0 All n p br 8, 26 A age i ae 40 Alph t b 6 Att n 40 A ici i 66, 6 , 62 A r 6 r t ll r 2 , 26, , 0 A pl t p p ll rn 2, 4 , 2 e i ae 6 , , 8, k, 06k, 24, 2 , 64, 86 A h 84: nd z , 44, , e i ae 6 k, 06k, 24, 64 84 r n l t 84 em i ii i 8 An t : f rb rn 2 , 84 b d n 2 t ph n n 84 t rb tt 0, 6 , 82, 84 l r p p n 84 A c y a si ae , 4 os ic i ae , 4 , 42, 44, 8k, 0 k, k, 8, 24, Andr ll p r 0, 66, 82 46, 86 An r t 4: h rp , 4, OS IC I O MIA 8 A o ii ae 6, , 4 , 42, 44, 02k, 0 , 08k, k, os ic i ae 4 , 24, 4 , 86 OS IC OI EA , 8k, 00k, 02k, 04k A o ii ae 42, 08k, 4 o i e i ae , 2, 0 k, 0k, k, , 86 An b : p n t t 42 r f rn 42 o i e i ae 2, A omma i ae 2, k, ac yce i ae 0, k, 6 An t 2: d d tr t 2, , r h p pl 46 Ant r t d 6 e i ae 6 , , k, 06k, 24, 2 , 6 , 86 A ici ae 6 , 64, 0 k, 0 k, , 60, 86 e i ae 0k, 06k, 6 A ici ae 6 , o i ae 48
88 r nt pr pl r l 48, 0, 6 C ysome i ae 2, , 6, , 4, 60, 66, , k, osci i 8 00k, 06, 2 , 2 , 62, 6 , 86 r n , 8k: th r 4, 6, 4 , C ysome i ae 66, 6 k, 62 r n ph l x r C YSOME OI EA 6 , 8k, k, 06k, 2 r h d v ll 6 , 62, 8 Chr p pl 6 uc i ae 6, 66, 6 k, 8, 00k, 06k, 62 Cici e i ae 2 r ll nt r t 84 Cici e i ae 2 k, 06k, 28 u es i ae 2, 6, , 44, k, 0 k, 20, 24, 42, Cici e i i 8 86 Cii ae 6, 6, 0 k, 0k, , , 86 pr t n Cii ae , pr t r l nt C ll n 46 U ES OI EA C z l nd , , , 8 y i ae 2, , , 40, 0k, 0 , 08k, 2 , 42, 86 C am i ae 2, 04k, 08k, 4 , 86 y i ae , 42 Cl b 2: d t 2, 4 , 2 Y OI EA , k, 00k, 04k, 06k, 0 k C A ICO IA 46 rr r pt : r h rt 6, 4 , C e i ae , 6, 44, 0 k, 0 k, 2k, 24, 48, 86 Cl r n 44 C ph t n rt 2 C E OI EA 4 , 02k C lp 2 Cl v n b l 2 , 2 , 68 C f l t r 28, , Cl v n n 8 C r n 2 , 26k, 2 Cocci u i ae 4 Ca a i ae 6, 8, 00k, 0 k, 0 k, 8, 24, 4 , C n ll : l n n 4, 2 , 4, 86 Cocci e i ae , 6, 4, , 04k, 0k, , 2 , Ca a i ae , 6, 2 , 60, k, 06k, , 4, , 26, 4, 86 26, 28, 2 , 8 , 86 Cocci e i ae 4, 4 Ca a i ae 2 , 22k, 28 Coe ome o i ae 60 Ca a i i 8 C l n 26: h rt l 26, CA A OI EA 20 Co o i ae 2 , 26k, C r b 2 Co y ii ae 2, , 6, 2, , 8, 04, 0 k, 0k, Ca io o i ae k, 20, 24, 6, 86 C rp l 2 , 6, 0 Co y ii ae , 6 Ca o i i ae 46 Co ym e i ae 22 C rp ph l 46 Co e a i ae 22 C th rt r pt l C pr t n 28 C t p l 26 C rt r Ca og a i ae 2, 6, 0, 0 k, 2k, 2 , 2, 86 Co ica ii ae , 6, 46, , 0 k, k, 4, 86 Ce am yci ae , 6, , , 44, 8, 6 , k, 06k, C rt r n 2 , 6 , 62, 86 Co y o i ae 4, 04k, 0k, 4, 86 Ce am yci ae 6 k, 6 Co y o i ae C r t n th : bb 2 p rr n 0, 8, , Cosso i ae 0, k, 66, 6 p l f r 2 C t l tr 8: z l nd 0, , 2 , 40, 2 C r d l 6 C t 6 : r p 6 , 60, 80 Ce y i ae 2 Cr ph l l t 6 Ce y o i ae 2, 0 k, 0k, , 86 C y a c i ae 46 Ce y o i ae Cr pt rph br v rn 48 Ch r d 60: tr h l d 6 , , 8 C y oce a i ae 66, 6 k, , 6 C ae osoma i ae 2, 44, 02k, 2k, 48, 86 Cr pt d n 2: br n 20 Ch t 44: r t d 44, 48, Cr pt l t 4 : f rr n 4 p ll d 4 Ch t d 44 C y o agi ae 6, 0, , 0 k, 0 , 0k, 2k, , Ch l dr 60: v r t 60, , 8 2, 86 C a co yi ae , 6, , 02k, 0 k, , 86 C y o agi ae 0, , 2 Ch l rt d 26 Cr pt ph 0 C e o a ii ae 4, , 6, 8k, 06k, 8, 4 , 86 C y o y c i i Ch l n r 6 C y osoma u i i C i oco i ae 4 Ct n r C o e i ae 2 , 26k Ct n n th n v z l nd 2 , 2 , 68 C o agi ae 68, 6 k, 64 Cucu i ae 4, 6, 48, 4 , 0 k, 0 k, 0 , 0 k, 2k, Chr l n : h p r 6 , 62, 8 dr n 6 20, 26, , 86
8 CUCU I O MIA 8, 42 Ep h r l n l 2 CUCU OI EA 46, k, 0 k, 02k, 04k, 2 Ep r 46 Cu cu io i ae , 6, , 2 , , 0, k, 06k, 6, E o y i ae 6, , 0 k, 2k, 20, 2, 86 8, 2 , 2 , 6 6 , 86 Euaes e i ae 2 , 28k, 08, 6 Cu cu io i ae 0, k, 66 E l p l f r 2 CU CU IO OI EA 4, 6 , 8k, 06k, 2 Euci e i ae 2, 02k, 08k, 4 , 8 C r 0 E n t t rt 2, 4 , 2 Cyc i i 8 EUCI E OI EA 00k, 02k, 04k, 08k C l x r 0, 0 , 0, 2: pr 4 , , 6 Euc emi ae 6, 00k, 0 k, 44, 8 p l t l 4 E l p br nn 6 , 2 , 6 , 8 Cyc a y i ae , 4 , E nn r l p 84 C ph n 2 E nn p r l p 84 Cy o i ae E d r 4 , 8: 4 , 24, 46, 4 C r t ph 6 : tr d n 6 , 64, 82 Eu e i ae , 4 , 0 , 46 Eug e i ae 64, 0 ac i ae 2, 2 Eumo i ae 66, 6 k, 6 ascy i ae E p n 28 t 4 : b n 4 , 4 , E pl t p 28 asy i ae 4 , 0 k, 2k Eu es i ae , asy i ae 4 , 4 Ex p n l 0 tr 6 Ex l p p t n t 62, ndr bl x rl 0, 8, Ex h 4 e o i i ae 24 e ico i ae , 44 Ga e uci ae 66, 6 k, 62, 6 r t 40: l t 4 , 2 Ga e uci i 66, 6 , 6 e mes i ae 6, , 40, 00k, 0 k, 0 , 08k, 2 , G d r : r l t 0, 8, , 84 th n 4, 46, 86 84 Moe au 84 e mes i ae 40, 08k Gn th r 6 e o o i ae 4, , 00k, 08k, 4 , 86 G n ph l 6 E O O OI EA 8, 00k Gr n r l r 4 , 4 , e o a i ae k Gym usi i 4 r l thr , 0 Gy i i ae , 4, 22, k, 06k, 0, 8 r pn d 6 : f ld 6 , , Gy i i ae 22, 0 ia e i ae 60 G r n : nv x l 22, 2 , 0, 68 h tt n 22 i o e i ae 4 , 46 G r ph n 2 n d r 4 : n t 4 , 46, 4 iscy i ae a oce i ae 2 , 2 k, 8, o ca omi ae 42 br r p ll r rn 2 , , 0 0 x z l r dr ph : p n p nn , t lb rp , t b r l t r r 6 : h tt 6 , 8, 84 yo o i ae 0, k dr br n 42, 4 , 4 yo i ae 4, 6 , k, 0 , 8, 42, 86 l 4: h l b 4 y as i ae 0, k, 40 p l p : pr l x , 2, 6 ysmo oce i ae 8, 4 a a i ae 2 , 22k, 2 n r pt p ll d 68, 64, 8 a a i i 8 y isci ae 22, k, 06k, 26, 2 , 0, 86 e o i ae y isci ae 22 t r x t t t 84 e e oce i ae , 0 k, 0 k, 4 , 8 E a e i ae , , , 00k, 0 k, 2 , 44 e e oce i ae , 4 E A E I O MIA t r r n v l nd , 4 , E a e i ae E E OME A 6 E A E OI EA 6, k, 00k, 0 k t r n h r t r , 40, 2 E mi ae 4, 6 , k, 08k, 42, 86 pp d 4 En r b ll 20 is e ae 2 , 8k, 0 k, , 8 E omyc i ae , , 0 k, 0k, k, , 4, 86 is e i ae 24 En v p , 4, h r 68 En hr tr t 2 , 0, 68 l p f l 84 E o ii ae 44, 48 o o a ameci ae , 0 k, k, 4
0 l p r , : t n 4, 4, th t n , 0 o e o i ae , 2 , 0 b r n th r l ph 4, 2 : l r 2 , 0, 68 b r : d pr , 2 n t n d r p 4, 42, n t r j b 6 dr n 24 o oca e i ae 4 , 44k, 2k, 4 y ae i ae 2, , 24, k, 08k, , 2, 8 r l 6 y ae i ae 24, x r n br d 2 , 26, 28, 68 y o i i ae , 4, 2 , 24, k, 08k, 8, 20, 0, uca i ae 6, 2 , 8k, 06k, 8, , 8, 8 , 8 uca i ae 2 , , 8, y o i i ae 2 , 0 yci ae , 6, , 00k, 0 k, 8, 24, 44, 8 Y O I OI EA 2 , k yc i ae 4 , 0 , , 46 y o o i ae 22, 2 , 0 t : br nn 4 , 46, 4 l n r 4 pl n ll l t t r , 6 , 8 4 l b : n b l , , 8 p ll 26 p r b : r n t 84 h tt n 84 p r 28 y a i ae 4, 0 k M r tr 6 : x l 6 , 60, 80 ph l 4, Mac a ii ae 6 , 60 p d n Mac au aci ae 6 p d n ll 2: r br p , , M r t l r t 0, 66, 8 Ma i i ae 8 Ill 4 M nd l t r ll 0, 6 , 82 In t p l n ll 26, 2 M r x r t I o e i ae 62, 6 , 00k, 02, 08, 0 k, M r n h n f r 84 M r p b r f rb rn 2 , , 28, 68, 84 aco so ii ae 2, , 0 k, 0 k, k, 4 , 8 M d 2 : ll n , n t l , tr x, br tt n , h l t n , t ll t ll , t ll l , t ll Ko y e i ae 44 b , t ll p ll , rv d n , d n n , d x, K h l dr : phr t 22, 0, 68 h tt , nt r t , l v p , l t r , n x, r , n t d , p v d , pl t , pr x , p l h ll , p n aemo oei ae 4 , 4 , 0 k, 0 k, 0 , 0 k, , t ll , n n , r x, tr t 84 8 M d n z l nd 2 , , ag ii ae 60, 6 , M l b : b f r t , b , lpt r t 84 r d 6 : br n 6 , 60, 80 M dr 2 , 84: nt r t 2 , 84 b l , ag ioi i ae 6 , 0 k, 60 p t , pr , f lt n , h pl p , Oma ama , amii ae 6 k, 6 v r n 84 am imi ae 2 , 8 Mega omi ae 40, 46 n t l n l t 26 Me a yi ae 2, , , 64, 02k, 0 k, 26, , 8 a gu ii ae , 0 k, 2k, 2 , 2, 86 Me a yi ae , a ai ae 4, 42 M l n phth l a ico ii ae , 4 Me asi ae 6, 44 rh n h b rb rn 0, 2 , 82 Me o o i ae 0, k, , 40 a i ii ae , 46, , 0 , Me y i ae 4 , 0 k, 2k, 4 , 8 a i ii ae , 4 M n 60 a i ii ae M r p th 24, , , 08 e ii i 8 M r p th z l nd 24, 2, 6 eio i ae 2, , 2 , 0 k, k, 8, 2, , 8 Me o ysii ae eio i ae 2 , 26k Me o ysii ae emo i ae 6 , 60 M t bl x 84 n x 46, 00, 0 , 04, : r nd 4, 46, 4 , 4 , M t rn l b 2 M t l r 64 p d pt r x 4 : n r p r 4 , 48, M t x n 44, 0 , : rn t 4 , 48, ici i i 8 M t phth l im ic i ae , k, 0 k, 4 , 8 M tr rrh n h rr t 8 im ic i ae , 08k, 4 M r br nt 4 n h : n r p , 4 , M r br nt l n t 4 , , 6 h r h tt n 4, 42, M r lph 4, 28: l t r 2 , , 6 d pl t 22, 2 , 68 Mic osi i ae 2 , 28k, 00k, 08, issomi ae Miga o i ae 2 , 22k, 28
Miga o i i 8 Orth p r 4, 0 M p 6 : l n t 6 , 8, p l 60 Or z ph l 48: r n n 48, , 6 p r ll l 84 b t t Oso ii ae 2 , 28k, 6 Mo omma i ae Oth 28 M n t 46, 00, 0 , 04, 0: p n ll 4 , 0, O y e i ae 2 , 28k, 6 Mo o omi ae 4, 46, 00k, 0 k, 04k, 0 , 0k, h r n 6 k, 4 , 0, 8 ae e i ae 2 , 2 k, Mo o omi ae 4 , 4 , 0 r l t n n n 84: r t l t 0, , M rd ll nt r t 8, 6, 8 Mo e i ae , 02k, 0 k, 6, 8 a amu i ae Mo e i ae 8, 6 r n h d n 2 , , 0 M t : l d bt r r n 4, , r t r h 28: l f r 84 fl x 84 Myce aei ae , 0k, r p r 24 Myce o agi ae 0, 6, 0 k, 0k, 20, , 8 r p lp l r 2 Myce o agi ae 6, rn d r t 4, 42, : l n l 4 M ll n 2 r p h r bd 66 M rt n z l nd 0, 66, 82 rt t n 6 MY O AGA , pr 44 e eca omi ae 8, 6 ace i ae 62 e i ae 4 a ose i ae 2 e ago ici i 8 d n , 20, 24, 42, 2 r pt 0: fr nt l , n t d l 84 tr n t , b l v rn 48 40, 2 r b 44: r f ll 44 r f p 44 e imy o i ae 60 r ph l 2 , 2: pr l n t 2 , 2, 6 a ac i ae , 4 , , 8k, 0 , 0k, 2k, , 8 emo yc i ae 2, 6, 6 , k, 06k, 2 , 6 , 8 a ac i ae 4 r 0: l t 0, 2 h l r 4 : n f r fr r l 0 h r pl x , 44, h l n r t 84 nd l t b r l t 2 , 28, 68 h l th r 2, b 6 , : tr d r 0, 2 , 6 , 82 oeoc a i ae 2 , 2 k, 4, 6 pt h pl p 84 hl n pr , 6 , 8 r ph nn t r n 84 oeos ic i ae , 4 , 0 k, 02k, 04k, 0 , 0k, i i u i ae 6, 46, 00k, 04k, 08, k, 8, 4 , 2k, 2 , 0, 8 8 hr t d 22 i i u i ae 46, 4 e a a i ae 0k oso e i ae , 40, 0 k, 0 , 08k, 4 , 8 h h r n p , , d ndr n : v t 40 z l nd 40, 4 , 4 ycoseci ae 2, 4 , 0 k, 2k, 4 , 8 th ld n p 84 h 4 : l b t 4 , 4 , th d r d nt : rl , 4 , 4 y o ae i ae 44 th t l 64: t t 64, 26, 6 , 80 h t ph 44: l br 24 v l 4 , 48, t l 4 Y O AGA 6 t pt n d l n 2 , 2, 6 h br x nfl t t t r h f rr n 2 , 6, 0 r t 0 ies i ae 2 , 28k, O l 28 i i a i ae 2, 62, 8, Oc e ii ae 24, 2 ime ii ae 60, 6 , O l nd nn r 84: l v l 84 i yo ii ae Od ntr v n , , l n t 4 Oe eme i ae 6 , 0 k, 0 k, 2 , 8, 8 l t p d 46: p r ll 46, 4 , Oe eme i ae 62, 8 l t 4, 48, 20, 26, , 6 O n h rt 8, 6 , 6 , 80 a y o i ae 0, k, 8, 06, 6 Ol t 28 l t p 4 : p l , 6 , 8 Oma ii ae 2, 4, 2 , 28k, 00k, d n l t p lp , 6 O l lb p nn 2 , , 6 O Y AGA 2, , , 8, 22, k, 0 k, , 2 , O d 6 22, 26 Onth ph r n l t , , rr t r f p nn 8, 24, 44, Or n l 84 r lph 02, 0 : b r 4 , 2 , 0, 6
2 iasi i ae 4 Sa i gi ae 6 , iasi i ae 4 k, 2k, 0 S lp n 6 : b l n t 6 , , io i ae 6 k, 6 S ph ph : n t 2, , 4 , 4 r n pl 8: r t l r 6 , 2 , 6 , 80 Sa i i ae 24, r t d r nt r t , 6, 8 S pr n 24 r d ntr 0: b l r t , v i e s , r nd 84 S pr t 0 l 0, , 40, 2, 84 d t , t , Sca i ii ae 26 28k, k, 26, p tr , r l , z l 84 Sca a aei ae , 6, , 0, 60, 8k, 06k, 2 , , os omi ae 6 , 04, 0 k, 0k, 8, 8 40, 8 o ei i ae 2 , 2 k, 00k, 4 SCA A AEI O M A 2 o e a e i ae Sca a aei ae 0, k, o o e i ae 4 , 44k, 2k SCA A AEOI EA , 6, 2 , 8k, 06k se a i ae Sca i i ae 2 , 22k, 2 se a i ae 2, , 6, 26 28k, 00k, 08k, , 4 S h r n 84 dh l p 6 Sci i ae 2, , , 0 k, 08k, , 4 , 8 d h l n r 6 SCI OI EA d phl h r tr l 2 , 4, 0 Sco y i ae 4 , 44, , 0, k, 8, 06, 6 seu o si ae 2 , 2 k, 6 S p d 2 d p rr 2 , 6, 0 Sc a i ae 64, 0 k, 0 k, 26, 60, 8 sy i i 8 Sc a ii ae 64, 6 e os ic i i 8 S r pt t 64 i ii ae 2, 6, 24, , 04k, 0 k, 2, 8 Scy mae i ae 2, , 26, 0 k, k, , 8 i ii ae 2 , 2 Scy mae i ae 26, i o ac y i ae , 00k, 0 k, 8, 4 , 8 Scym i ae 4 i i ae 6, 42, 02, k, 24, 4 S n 4, t n : p 42, , 24, 4 , 4 t t 42 S p d ph l 2 , 28, 4, 0 yc ome i ae , 0k Se ico e i ae , 4, n r ph r 24 Sh rp v n t 68, 64, 8 y oc oi ae 62, 0 k, 0 k, 2 , 8, , 8 Si i ae 2 r n t f t v 0 S lph t l 08: n t d 2 , 4, 0 y i ae 62 Si a i ae 6, 4 , 0 k, 02k, 0 k, 0, , 8 Si a i ae 48, Q d 28 S lv n 48 S rrh 60 v p l 40 S t n d d 2 h rdt 24: p d tr x 24, , 6 S r n 46 nt n d t v ntr 4 , 26, 4 , 4 S ae i ii ae 2 , e o ii ae 4 , 44k, 2k, 26, 4 Sph r th r x 2 e o i i 4 S o y i ae 6 k i o y c i ae 68, 6 S a y i i ae 2 4, 6, 26, 00k, 0 k, 08k, k, h n rh n h r f l 68, 2 , 6 , 8 6, , 8, 26, 2 , , 8 i i o i ae 8, 8k, 00k, 0 k, 8, 26, 6, 8 S A Y I O MIA 2 h p t n 8: l br 8, 6, 8 S a y i i ae 2 , 2 k, h z n nt , 6, 8 S A Y I OI EA 2, , 24, k, 00k, 02k, 04k h z p rth 4 : d n 4 St b p n 42 i o agi ae 04 St ph n rh n h n l t 84: l n 0, 66, 8 Y C O O A 6 St th p 0: nv x 84 y c o o i ae 0, St l n b nt r t 2, 4 yso i ae 20, k, 06k, 8, 28, 8 Sy ca y i ae yso i i 20 Sy esi ae 2 h z b 4 S rth n l r 2 h z d t pr pr 20, 28, 68 S rph t d , 84: r n t , , 8 d l 4, d n rt 20: t b l 2 , , 6 ac y o i ae 2 , 2 k, 4 t n t 8 n r n p n 84 n h l 6 : ph r 26 S l 28: l n t 2 , 4, 0 a sos e i ae 44 Sa i gi ae 62, 00k, 02k, 08, 0 k, 0k, , 8 h : n l r 62, 8, t l ph r d 2
h d 62 n br 6 e e io i ae , 4, 6, 6, , 60, 8, 0 k, 0 k, 0k, , , , 8, 8 e e io i ae 60, 6 , 8 E E IO OI EA 2, , 6, 8k, 00k, 0 k, 04k n br d r t n 4 h ll p l t 2, 2, a oc e i ae 44, 48 h l ph l n t 62, 8, h r : f ld , 44, h rt v l , 2, 6 h t h ll 84 n l t p 24, , 6 ec i ae 2 , 22k, 2 ec i i 8 i a i ae 24 r b l 6 r h l d 40 ic e o i i 40 r h lp n 6 i o i ae 40 r ph ll 6: h p d ll 6, , ogi ae , 0, 8k, 06k, 8, 8, 8 r d r 40: t 4 , 46, 4 ogossi i ae , 6, 4 , 04k, 2k, 26, 4 , 48, 8 ogossi i ae 4 k, 2k, 48 r x b r 0, 8, h n p t b r 84 b p pt v l 2 ph t r r 20
U o i ae , 60, 0 , 02k, 0 k, , , 8 Ul 60 Ul t p 60
r n t 2, : tr t ll 2, 4 , 2
n n p n p 6 , 8 e osce i ae 2 l h 6 y o i i ae 64 l ph l 64: n t d 64, 60, 80 l t l t p nn 6 , 6 , 80
r t d 26: l f r 26 ll n : l r 2 , 2 , 68 t ll rd 84 n d l 0: h th n 0, 2, 6 d bl t n 0 p l 84 o i i 8 o o i i ae 60 l d n 60 d 6 o e i ae , 02, 0 Ζ r n 6 , 6 , 80
4 O ICES GA A UI is se ies o e e ee occasio a u ica io s as Kua w aka ū ia ē ei ā a gi uka uka ei ee es a is e o e cou age ose wi e e w aka au au ki ga o u ga w ai mā au a ga k ow e ge o u is co cise ye com e e si e kia w akā u i ga mea e ā a a ki ga ka a e e o
accou s o e eme s i e ew ea a au a. ' iu i e i. e ā ua o u ga ē ei ā a gi e se ies is o essio a i i s co ce io a uka uka, e ga i, ko e ia ia kia mā ama ai e ese a io , ye e e y e o is ma e o o i e u i u i, kia mō io ai e ma ia ki ga o u o ia esou ces o i e i ica io a i o ma io a gā a a, o ia gā a a, ā, kia w akā i i e mā au a e accessi e o e o s ecia is . a ga e ā a a ki a ā ou. n f . ea s wi o ma i e i e e a es Κo ē ei uka uka n f l nd kāo e e o y, si ce e e e a es a e we ocume e , ā a a ki ga ka a e e moa a, a ā, ki ga ika, ki a ma i e o ms a e co e e y e se ies M r n ga mā ai ai ā ei. E i o mō io ia a a ga n f . . ka a e e. Kei o o i ga uka uka e kīa a a M r n n f l nd n u i u e ā a a ki ga ka a e e moa a ga ika me ga mā ai ai, oki. Co i u io s a e i i e om a y e so wi e equisi e s ecia is ski s a esou ces. Ma e ia om e . . A o o Co ec io is a ai a e u i u i ga. Ko e o o ki ga o u ga kia ukua o s u y. maiāko iko i a a kou ou uka uka. e E wā ea ga ka a e e e kīa a a ko e ew ea a A o Co i u o s s ou iscuss ei i e io s wi a o Co ec io ei mā aki aki māu. mem e o e I e e a e Sys ema ics A iso y G ou o wi e Se ies E i o e o e comme c Me w ākī ō kou ou w akā o ki e mema o e kā ui i g wo k a ecessa y gui a ce wi e gi e . o u o u o n ika a a, ki e E i a ā ei, i mua i e īma a ga u i u i.
Su sc i e s s ou a ess i qui ies o n f . ., Ma aaki W e ua ess, a ca e e ga kai oko uka uka. Me u i ki e n f sea c , .O. o 40, i co 8 2, ew ea a . . ., Ma aaki W e ua ess, a ca e e sea c , .O. o 40, i co 8 2, ew ea a . Su sc i io ca ego ies: A — s a i g o e s a i oice wi e se wi eac ew issue, as soo E ua ga ūmomo kai oko: A — Kai oko ūmau a e u ica io as ossi e — omo io a ka ukua a uka uka, ia uka uka, me e kau e, i ie s wi o e o ms wi e se om ime o ime. mu i o u i e ā ga o aua uka uka. — ka ukua ga ā ui a ake, a ō a wā, a ō a wā. e ai ices (see i es i i , age 8 i c u e ackagi g a su ace os age. Su sc i e s i e u u ( i o ia e w ā a gi 8 : Κo e kō aki a ga ew ea a a Aus a ia ay e i ica e me e a e kui i kei o o i e u u. Me u u kou ou e amou i $ GS is i c u e i e ice. O e o o a a i iu i e i me Ā i e ei ia ki ga ā a o su sc i e s ay e is e ice i $US, o i s iu i e i. Ko ē a i a u me u u e w aka e ga equi a e . ga ā a Ma ike a. ack issues o a um e s a e a ai a e, a ew E oe a a ga uka uka o mua. Me emea e ia ia su sc i e s wis i g o o ai a u se o a se ec a a koe ki e ka oa o ga uka uka, o oa kiamai io may eques a iscou . ookse e s a eke i o e u u. E ekau ai ē e i e eke i o o e su sc i io age s a e o e e a a e iscou o u u ki ga oa oko uka uka. e e ce .
— — 2° ° 4° ° 6° ° 8° 2° ° 4° ° 6° ° 8°
° °
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WO Waika o Three Kings ❑ 42° Ι 42°
❑ 40° 40° C a ams Sou Is a S a es ❑ u e ❑ ι S ou y 4 ° CO Ce a O ago 4 ° W ππιτ Λι Λ.. Λ u e i A i o es ❑ ΙΛιιιιιΙΟι[ aιιππuιπι, io a Auck a s ❑ KA Kaikou a 42° Cam e ❑ ΑΙΙΙΜΙΛιιιΛ M Ma o oug ΑΙΙΙΒ ΙΙΙΙΙΙιι MC Mi Ca e u y ΙΙΙΙΙΙΙΙΙΜΒ 0 -420 _ΙΙΙΙΙΙΙΙΙΙΙΙΙι MK Macke ie ι.ιιιιι·ιι.·ι . 4 ° 4 ° Ι ΙΙΙ ιιΛ ι.ιι·Λιι .ι ϋιί ίπ ι ww, ι W ιίιιιιιπίιιuιππΜι ιί 44° ΙΙΙΙΙιιΛ.ι ιιιΙιιι" 44° ΙΜΙΙ.··ΙΙ·ι···ΙΙΙι λιΙ.ΙΙΙW ιΛΛιι Ι, :i•iiiii aiiiΛiiiii Αιιιιιιιιιιιιι•ιιιιιιΙ ιιιιιιιιιιιιιιιιΛιιιι ι.]ιiιιιιιιιιιιιΛιιιιιΙ , 4 ° 4 ° : ΛΛ uΛιιΙΙΙΙΙριιιΓ ■4 ΙΙιΙΙιΙΙΙΙΙΙιιΙΙΙΙ=ιι ΩιιθιιιιιΛιιιΙιιι ιΙ C o Ca e u y ϊ.ϋΛιι•ιιιι ιιιιιιιιι ΩιιιΙΜΙΙΙιΛ ΙΙΙΙ ΝΝ e so 46° λι.ιι ιιι.ιιιιιιιιιιι 46° O O ago akes SC Sou Ca e u y S Ma o oug Sou s iι•ii 4 ° 4 ° SI S ewa Is a S Sou a W Wes a 6 ° 68° 6 ° 0° ° 2° ° 4° 6 ° 68° 6 ° 0° ° 2° ° 4°
A ea co es a ou a ies use o ca ego ise ase ma o o g co ec io oca i ies is s ecime oca i y a a (a e C os y t l 6 may e o oco ie wi ou co y ig e ease
— 6— o o k s a 0°S Ke ma ec Is a s
• o owe Is a SOU ee Ki gs s a s ΡΑ CI IC ° ASMA Ο C ΕΑ SEA O IS A
40°
SCA E (km a 4 °S a i u e C a am Is a s o 0 200 400 SOU 4 ° 1 ι ι ι ι IS A
S ewa ou y e S a es • Is a Is a s
A i o es , 0° s a s Auck a s a s
Cam e Is a E EW EA A SU EG O (e c u es o owe, o o k, a Macqua ie is a s e ce i e co e o e a imi a oogeog a y Macqua ie Is a °S 60°E 6 ° 0° ° 80° °W
— — I ES I I U A AI A A AA e e a ia (I sec a: ysa o e a • r n A. M nd & Ann tt K. W l r IS 0 4 0668 • 2 ec 82 • 20 . $2 . 2 Oso ii ae (I sec a: Co eo e a: S a y i i ae • . l n M C ll IS 0 4 06688 • 2 ec 82 • 6 . $ 8.60 A i i ae (I sec a: Co eo e a • .A. ll IS 0 4 06 0 4 • 2 ec 82 • 2 2 . $4 .00 4 E io yoi ea e ce E io yi ae (A ac i a: Aca i • .C.M. M n n IS 0 4 06 4 • 2 o 84 • 44 ρ. $2 . E io yi ae (A ac i a: Aca i: E io yoi ea • .C.M. M n n IS 0 4 06 46 8 • 4 o 84 • 28 . $2 . 6 y ae i ae (I sec a: Co eo e a • .G. Ord h IS 0 4 06 4 6 • 2 o 84 • 64 . $ 8.60 C y os igma a (A ac i a: Aca i — a co cise e iew • M. xt n IS 0 4 06 62 • 8 ec 8 • 2 . $2 . 8 Ca i o i ae (I sec a: i e a • . r
IS 0 4 06 64 6 • 24 e 86 • 88 . $ 8.60 o u a (I sec a • S. . x n
IS 0 4 06 6 4 • 24 e 86 • 2 . $ 8.60 0 u u i e a (I sec a: ysa o e a • r n A. M nd & Ann tt K. W l r
IS 0 4 06 84 0 • 22 Se 86 • 44 . $ 4.6 seu ococci ae (I sec a: emi e a • .M. C x
IS 0 4 06 • A 8 • 2 2 . $4 . 2 om i i ae (I sec a: yme o e a • A.C. rr IS 0 4 02 0 • o 8 • 60 ρ. $ . E cy ae (I sec a: yme o e a • .S. IS 0 4 02 • May 88 • 2 . $44.
4 e i o e a — a o a e ca a ogue, a keys o ami y g ou a a . S. d l • IS 0 4 02 8 8 • 2 Se 88 • 264 . $4 . Am osi i ae (I sec a: yme o e a: ia ii ae • I. . nn IS 0 4 02 8 • 0 ec 88 • 68 . $ . 6 e icu i ae (I sec a: e i o e a • n nn r & Chrl t ph r W l n n
IS 0 4 02 8 2 • 28 A 8 • 2 . $22. Myma i ae (I sec a: yme o e a • .S. & E.W l nt n IS 0 4 02 42 0 • 28 A 8 • 00 . $24. 8 C a ci oi ea (I sec a: yme o e a — i o uc io , a e iew o sma e ami ies .S. & Ε.W. l nt n • IS 0 4 02 4 • 2 Aug 8 • 6 $24.
— 8— Ma o ea (I sec a , wi a e iew o as ec s o u c io a mo o ogy a io ogy • G.W. • IS 0 4 02 8 • u 0 • 6 $24. 20 i io i ae (I sec a: i e a • A. rr n IS 0 4 02 • o 0 • 28 . $ 4. 2 Ma ga o i ae (I sec a: emi e a • C. . M r l IS 0 4 0260 • 2 May • 24 . $ 4. 22 o o emou i ae (I sec a: eco e a • I. . M ll n IS 0 4 02 8 8 • 2 May • 64 . $24. 2 Scia o i ae, Me e e i ae (I sec a: i e a wi a ge e ic e iew o e o ic o o i ae • . . l • IS 0 4 0262 • a 2 • 4 ρρ $2 . 24 e e i ae (I sec a: i e a • . n b r IS 0 4 026 2 • 4 Ma 2 • 40 $ 4. 2 Ce co i ae (I sec a: omo e a • K.G.A. lt n & C. . M r l
IS 0 4 026 6 2 • 2 May 2 • 40 . $ . 26 e e io i ae (I sec a: Co eo e a : ca a ogue o y es a keys o a a .C. W tt • IS 0 4 026 • u 2 • 0 . $2 . 2 A a c o e i ae (I sec a: eco e a • I. . M ll n
IS 0 4 0 644 8 • 8 e • 0 . $2 . 28 a ae o Cu cu io oi ea (I sec a: Co eo e a : a sys ema ic o e iew r nd M. M • IS 0 4 8 04 0 0 • 4 u • 226 . $ .00 2 C y o y c ae (I sec a: Co eo e a: Cu cu io i ae
C. .C. l • IS 0 4 8 04 8 2 • 2 ec • 08 . $6 .00 0 e ia i ae (I sec a: e i o e a • .S. d l
IS 0 4 8 04 24 • Ma 4 • 64 . $42. 0 a i i ae (C us acea: Am i o s • K.W. n n S 0 4 8 04 6 • Oc 4 • 28 . $ 6.00 2 S eci ae (I sec a: yme o e a • A.C. rr IS 0 4 8 04 4 4 • Oc 4 • 2 . $ . 0 Mo a i i i (I sec a: yme o e a • .A. rr IS 0 4 8 04 8 • 8 May • 82 . $2 . 4 A ici ae (I sec a: Co eo e a • .G. W rn r& .S. Ch ndl r
IS 0 4 8 04 4 6 • 2 u • 64 . $26. 0 Cy i ae, Aca osoma i ae, a e a omi ae (I sec a: e e o e a :
sys ema ics, geog a ica is i u io , a ioeco ogy • M. C. r v èr IS 0 4 8 0 0 2 • 2 o • 2 . $42. 0 6 e o e ii ae (I sec a: E eme o e a • . . n & W. . t r IS 0 4 8 0 0 • Aug 6 • 44 . $ . 0 Co eo e a: ami y g ou e iew a keys o i e i ica io • . Kl z & .C. W tt • IS 0 4 8 0 2 8 • u • . $4 . 0
— — n f l nd K t A t n p A t r
b r
4. oue e iew a
Kl z & C W tt with illustrations by D.W. Helmore O U A SUMMA Y
C ECK IS O A A
Ν .
E IEW O AMI IES
KEYS O A A
. i A O OMIC I E This is a PDF facsimile of the printed publication, and is fully searchable. It is supplied for individual use only and is not to be posted on websites (links should be made to the page from which it was downloaded).
ρ rt f th r v r d b p r ht b r pr d d r p d n n f r or by any means (graphic, electronic, or mechanical, including photocopying, recording, taping, information retrieval systems, or otherwise) without the written permission of the publisher.
Fauna of New Zealand website copy 2010, fnz nd r r h nz
Klimaszewski, J.; Watt, J.C. 1997: Coleoptera: family-group review and keys to identification. Fauna of New Zealand 37, 199 pp.
Date of publication: 13 August 1997
Fauna of New Zealand, ISSN 0111-5383; 37 ISBN 0-478-09312-8
New Zealand Coleoptera. Scanned images provided by Stephen Pawson (www.bugz.org.nz). OCRed text corrected for this searchable PDF by Trevor Crosby, FNZ series editor, 6 July 2010. Users may extract text from this PDF for their own use, but must check it against the original document for text sequence, accuracy, and formatting.