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Memoirs of the Museum of Victoria 56(2):659-666 (1997) 28 February 1997 https://doi.org/10.24199/j.mmv.1997.56.67

BIODIVERSITY OF NEW ZEALAND (INSECTA, COLEOPTERA)

J. KLIMASZEWSK.I

Manaaki Whenua — Landcare Research, Private Bag 92170, Auckland, New Zealand Present address: BC Research. 3650 Weshrook Mall, Vancouver V6S SLS, Canada

Abstract

Klimaszewski, J., 1 997. of New Zealand beetles (Insecta: Coleoptera). Memoirs of the Museum of Victoria 56(2): 659-666. Approximately 5235 are described for New Zealand, including 354 introduced. They belong to 82 families in two suborders, and . The New Zealand fauna is distinguished by the absence of many major lineages, a high level of endem- ism. which in many groups is over 90% at the specific level and over 43% at the generic level (e.g.. Staphylinidae), and the radiation of many groups of genera and species. The origins of New Zealand's beetle fauna are still poorly understood. They are likely to be varied, includ- ing Gondwanan elements and elements which arrived here by short and long-distance dispersal recently and in the remote past. The size of the New Zealand beetle fauna is con- sistent with species number/land area relationships in other areas around the world.

Introduction Zealand beetles is that of Kuschel (1990), in the suburb of Lynfield, Auckland, in which 982 The beetles are the largest order of organisms, beetle species were recorded in a diverse veg- with over 350 000 described species world- etation including remnant forest, pastureland, wide. and suburban garden. There are approximately 4881 described Factors responsible for the diversity and native and 354 introduced species of beetles in nature of the present-day New Zealand beetle New Zealand (Appendix 1), representing c. 1094 fauna are: genera in 82 families (Klimaszewski and Watt, 1. Gondwanan origin of some of our biota; in press). The total number of species is esti- 2. approximately 80 million years of geographic mated at around 10 000 to 10 500 (Watt, 1976: isolation of New Zealand, which has resulted Kuschel, 1990). In comparison, the vascular in a high level of generic endemism and plant flora includes about 2500 species, and the approximately 90% endemism at the specific terrestrial vertebrate fauna about 350 species level; (Watt, 1976). Watt (1976) estimated at 20 000 3. changing climate, changing shorelines (Oligo- of described and undescribed the number cene bottleneck) (Cooper and Millner, 1993; species of terrestrial and freshwater Arthropoda Cooper and Cooper, 1 995), orogenies, glacia- probably occurring in New Zealand. tion, and volcanic activity; and Zealand beetles belong to two suborders, New 4. absence of terrestrial mammals and other Polyphaga. and Adephaga and and plants which have dominated in , are absent , present ecosystems elsewhere in the world. from New Zealand. Four families (Archeocryp- and ) are ticidae, Gyrinidae, , Origin of New Zealand beetles present only as introduced species. One, Chalco- Zealand region its ancestral biota dryidae, is endemic. Eleven families are rep- The New and resented by more than 100 species: Curculioni- originated in the fragmentation of part of an )" including Pselaphinae ancient supercontinent, (Cooper and dae ( 1 32 1 Staphylinidae 1 and Cooper, 1 In the (1021), Carabidae (445), Colydiidae (196), Cer- Millner, 993; Cooper 995). 80-85 (million years ambycidae (188), Scydmaenidae (165), Chry- late c. mya Elateridae ago), the New Zealand region had separated somelidae ( 1 56), (144),

Scirtidae ( 1 25). from the remainder of Gondwana through (135), Tenebrionidae ( 1 34), and broadening Some smaller families — , , active sea-floor spreading and of the Tasman Sea. The New Zealand region with its Hvdraenidae, Corticariidae ( = ), has been geographically isolated for Le'iodidae, , and — biota thus approximately the last 80 mya, and during this are also well represented in New Zealand. The most comprehensive local survey of New period has undergone an active tectonic and

659 660 .1. KI.IMASZEWSKI

Figures 1-4. I Horelophus mlkeri beetle . Qrchymont, ofthe endemic subfamily Horclophinae (Hydrophilidac): litorea 2, Microsiipha Broun, endemic species of Mierosilphinae (Staphylinidae): 3, Ampkctopus palticorni's Broun (); 4. famida agrestis Broun, endemic species (). BIODIVERSITY OF NI-AV ZEALAND BEETLES (,(.1

of ; 6, Nothoderodonm Sharp, only New Zealand member 5-7. 5, Hrouma thaiacica Figures minimis Sharp (Jacobsonudae). 7, aSaphophagus gourlayi Crowson (); 662 J. KI.IMASZEWSKI

volcanic history. These events have had pro- tralia, and South America. Some southern tem- found effects on the composition and distri- perate from families Hydraeni- bution of plants and animals, including beetles, dae, , , , Pselaphi- and influenced their . Certainly some dae, and Staphylinidae show 'transaustral of the contemporary New Zealand beetles have disjunctions' (occur on two or more widely sep- evolved from ancient Gondwanan forms. The arated southern land areas) — New Zealand, modern beetle fauna consists of ancient, vari- Australia, South , and/or South Amer- ously changed lineages, elements introduced by ica. dispersal over short and long distance (e.g., Comparison of Staphylinidae in New Zealand groups dispersed passively over water as a result and other areas of the strong westerly winds and currents around the 40th parallel), and species intentionally and The Staphylinidae in New Zealand accidentally introduced. The fauna was greatly includes approximately 936 native and 85 influenced in the last 1000 years by people who adventive species (Klimaszewski et al.. 1996. brought devastation to native forest and intro- A comparison of the estimated numbers of duced harmful exotic animals — possum, rats, species of rove beetles in New Zealand, the goats, deer and others (Ramsay, 1978). Hawaiian Islands, Florida, the British Isles, Cen- Support of the hypothesis that many of our tral Europe, Fennoscandia, Australia, and North beetles are of ancient origin comes from the high America north of Mexico is presented in Table endemism, over 90% at the specific level, with 1 . Species numbers are the result of many factors several endemic subfamilies and tribes and besides land area, including latitude, climate, many endemic genera (Watt, 1982). Several topography (resulting in diversity), and usually small families are represented in New the geological and biological history of an area Zealand by diverse endemic forms (e.g., Byrrhi- (including the impact of Pleistocene glaciations dae, Colydiidae, Hydraemdac. Melandryidae, and the degree and duration of isolation from Ptiliidae, Scirtidae and Scydmaenidae). Native other areas). species are those primarily associated with the New Zealand's staphylinid fauna is compar- lowland forest which has prevailed throughout able in size, in relation to land area, to the faunas most of New Zealand's geological history, the of a diversity of areas. This is quite striking in alpine , the tussock grasslands, and the view of New Zealand's much longer isolation subantarctic islands (Kuschel. 1990). Intro- from other lands (compared to the other areas in duced species are mostly associated with anthro- Table 1) and its perhaps consequent lack of sev- pogenic habitats (Kuschel, 1990). Many groups' eral major lineages of Staphylinidae. The distribution patterns are suggestive of a tremendous topographic, climatic, and veg- Gondwanan or pre-Gondwanan origin: for etational diversity of New Zealand is likely to instance, the remnant relict genera of tenebrion- have enhanced speciation. There could also be a oid Pilipalpinae persisting in Madagascar, New connection between the absence of several lin- Zealand, southern South America, and Aus- eages and the extensive New Zealand radiations tralia. The occur in New Zeal- in Microsilpha, Sagola. *Euplectopsis, Eupines, and, Australia, southern Africa, Madagascar, Sepedophilus, O/igota, *Paratorchus 1 Hype- and the Holarctic region. The staphylinid Stylo- romma. Othius, and 'Quedius' [*endemic gen- gymnusa subantarctica Hammond is known era]. Far more knowledge of the ecological roles only from the Auckland Islands, with its closest of staphylinid groups both in New Zealand and relatives exclusively Holarctic. The New Zeal- elsewhere is needed however, to test such an and jacobsoniid beetle Saphophagus minutus hypothesis. Sharp was considered by Crowson ( 1 959) as a The origins of New Zealand's staphylinid relict coeval with the tuatara. The Chaetosoma- fauna are still poorly understood. They are prob- tidae are known only from New Zealand and ably varied, since the fauna includes over 20 Madagascar, and the Phycosecidae only from genera or higher groups shared only with other New Zealand and Australia. The small family southern temperate regions (Newton, 1985). as Cavognathidae, with species occurring in birds' well as numerous more widespread lineages nests, are also known only from New Zealand (e.g., Sepedophihis, Gyrophaena. Oligota, Myl- and Australia. The New Zealand laena, Polylobus, Bledius, Carpelimus, Oxytelus, are closest to species from Chile. Omaliine sta- Lathrobiina sensu lato, Othius, Quediina, phylinids of the Metacorneolabium (22 Caftus, and Philonthm). Phylogenetic studies of species) are distributed in New Zealand, Aus- all lineages within and outside New Zealand are BIODIVERSITY OF NEW ZEALAND BEETLES 663

Table 1. Comparison of land areas and estimated numbers of speeies of Staphylinidae (including Pselaphinae, , Dasycerinae, and other groups) for several parts of the world (Klimaszewski, 1996).

Zoogeographical area Land area Estimated Source of estimate 2 (km ) species

USA.: Hawaiian 16 700 100 Nishida(1994) Islands USA.: Florida 140 800 498 Peck and Thomas, unpublished checklist New Zealand 268 800 936 Newton and Thayer (1995), unpublished checklist; Chandler (1991), (Pselaphinae)

British Isles 3 1 5 000 1046 Pope (1977) Central Europe 882 250 1949 Lucht (1987); Lohse and Lucht (1989) (Austria, Czech Republic, Germany, Poland, Slovakia) Fennoscandia 1073 500 1222 Silfverberg(1992) (Finland, Norway, Sweden) unpublished checklist; Australia 7 692 300 2441 Newton & Thayer (1995), Chandler, in litt. (Pselaphinae) (Pselaphinae) 19 115 250 3974 Arnett (1985); Chandler (1994), (N of Mexico)

Chandler, D.S., 1991 and 1994. Unpublished checklist needed in order to understand the history of the of Pselaphinae (Staphylinidae). New Zealand staphylinid fauna. Cooper, A. and Cooper, R.A., 1995. The Oligoecne of species for Australia and Estimated number bottleneck and New Zealand biota: genetic record known described and New Zealand includes of a past environmental crisis. Proceedings of the adventive undescribed species, but excludes Roval Society of London li 261: 293-302. species. Adventive species are also excluded Cooper, R.A. and Millner, P.R., 1993. The New and new from the Hawaiian Islands number, but not Zealand biota: historical background ecology and evolution 8: 429- from the other geographic areas, where the num- research. Trends in 433. ber of adventive species is not known, but prob- Crowson, R.A., 1959. Studies on the Dermestoidca ably very low. Known but undescribed species (Coleoptera), with special reference to the New included for Florida, but not for the are also Zealand fauna. Transactions ofthe Royal Entomo- where the number is not remaining areas, logical Society of London 3(4): 81-94. low. of known, but probably Klimaszcwski, .1. and Watt, J. C, in press. Beetles New Zealand: family-group review and keys to Acknowledgements identification (lnsecta: Coleoptera). Fauna of New Zealand. and W.M. Beever, C.T. Duval, M., 1 996. I thank R.E. Klimaszcwski, J., Newton, A.F . and Thayer, Auckland, for Wouts, all of Landcare Research, A review of the New Zealand rove beetles version of their useful comments on the original (Coleoptera: Staphylinidae). New Zealand were pro- Zoology 23: 143-160. the manuscript. Funds for the research Journal of Science G., 1990. Beetles in a suburban environment: vided by the Foundation for Research, Kuschel, a New Zealand case study. DSIR Plant Protection and Technology under contract no. C09308. Report 3. 119 pp. Lawrence, J. F., and Newton, A. F., 1995. Families and References subfamilies of Coleoptera (with selected genera, family-group handbook o] notes, references and data on R.H., 1985. American . A Arnett 779-1006 in: Pakaluk, J. and Mexico. Gaines- names). Pp. the insects of America north of S.A. (eds), Biology, phytogeny, and Publications. 850 pp. Slipinski, ville, Flora and Fauna 664 J. KI.1MASZLWSKI

classification oft 'oleoptera: papers celebrating the Nishida, Ci.M. (ed.), 1 994. Hawaiian terrestrial arthro- ( 80th birthday ofRoy 'rowson. Mu/.eum i Instytut pod checklist, second edition. Bishop Museum I'AN: Zoologii Warsaw. technical Report 4. iv + 2X7 pp.

Lohsc, (i.A. and W.I I., 1 Lucht, 989. Die Kafir Peck, S.B. and Thomas, M.C. (unpublished). A list of Mitteleuropas; l. Supplementband mit Katalog- ( oleoptera of Florida. teil. Goecke and livers: Krefeld. 346 pp. Pope, R.D., 1977. Coleoptera and Strepsiptera. In: Lucht, W.M., 19X7. Die Kdfer Mitteleuropas; Katalog. Kloct, G.S. and Hincks, W.D. (eds), A cheek list of Goecke and Evers: Krefeld. 342 pp. British insects (second edition). Handbooks for Newton, A.I-'., 19X5. South temperate Staphylinoidca identification of British Insects 11(3). Royal (Coleoptera): their potential for biogeographic Entomological Society of London: London, xiv + analysis of austral disjunctions. Pp. IXO-220 in: 1 05 pp. Hall, (i.H. (cd.), , phytogeny and Ramsay, G.W., I97X. A review of the effect of rodents zoogeography of beetles and . Dr W. Junk on the New Zealand invertebrate fauna. New Publishers: Dordrecht. Zealand Lands and Survey Department, Newton, a. I-', and Thayer, M.K., 1995. Protopselaphi- Information Series 4: 89-97. nae new subfamily for Protopselaphus new genus Silfverberg, H., 1992. Bnumeratio Coleopterorum From Malaysia, with a phylogenetie analysis and l-'ennoseandiae, Daniae el lialliae. Hclsingin review of the group of Staphylinidae Hyonteisvaihtoyhdistys: Helsinki, v + 94 pp. including Pselaphidae (Coleoptera). Pp. 219-320 Watt, J.C., I 976. A biological survey in New Zealand.

in: Pakaluk, .1. Slipmski, and S.A. (ed.). Biology, The New Zealand Entomologist 6(2): 138-143. phytogeny and classification ofi 'oleoptera: papers Watt, J. C, 19X2. 1981 Presidential address: New Zeal- celebrating the 80th birthday of Roy A. ( 'rowson. and beetles. The New Zealand Entomologist 7(3):

Mu/.eum i Instytut Zoologii I'AN: Warsaw. 213-221.

Appendix I Inventory of New Zealand beetles. Classification adopted from Lawrence and Newton (1995). Suborders and families with over hundred species are in bold.

Taxon Approximate number of Approximate number of recorded native species introduced and adventive species

Suborder ADKPHAGA Siipcrfamily CARABOIDEA 1. 6 2. Carabidae (including Cicindelinae) 426 19 3. Dytiscidac 12 1 4. CJyrinidae 1 1 Suborder POLYPHAGA Series STAPHYL1NIFORMIA Superfamily 5. 70 5 6. 22 6 Superfamily STAPH YLINOIDEA 7. Hydraenidac 32 8. Ptiliidae 48 8-9 9. Agyrtidae 2 10. Lciodidac 112 1

1 1 . Scydmaenidae 201 1 12 Staphylinidae 936 85 Scries SCJRTIFORMIA Superfamily 13. Scirtidae 125 (i 14. I 15. 8 2 .

BIODIVERSITY OF Nl W ZEAI AND mill is 665

Appendix l Continued.

Taxon Approximate number of Approximate number of recorded native species introduced and adventive species

Scries SCARABAII1 ORMIA Superfamily 16. Lucanidae 24 4

1 7. Trogidae 1

1 8. Scarabaeidae 132 12 Series ELATHRll ORMIA Superfamily 19. Superfamilv BYRRHOIDIA 20. Byrrhidae 79

2 1 Dryopidae 4 22. 16

23. Limnichidae 7 1

I) 24. 1 25. 6-8

26. Chelonariidae I Superfamily 27. 22 28. Elateridae 132 3

29. Lycidae 1 30. Cantharidae 40 1 Series Superfamily DERODONTOIDEA 31. Derodontidae Superfamily 32. 3 33. Nosodendridae 2 34. II 6 7 35. (incl. Lyetinae) I 36. Anobiidae (incl. Ptininae) 28 U Series Superfamily 37. Trogossitidae 24 I 38. Chaetosomatidae 4 37 2 39. ( leridae 40. Phycosecidae 1 41. Dasytidae ^^ Superfamily II 42. Nitidulidae 21

I 5 43. 44. Phloeostichidae 2 7 5 45. 46. 1 47. Laemophloidae 3 ! 48. 3 I 5 o 49. Cavognathidae 23 12 50. 8 51. Languridae 9 52. 6 1 53. 51

666 J. KLIMASZEWSKI

Appendix 1 Continued.

Taxon Approximate number of Approximate number of recorded native species introduced and adventive species

54. 5 55. 6 1 56. 22 18 57. Corylophidae 19 2

58. Corticariidae ( = Lathridiidae) 53 11 Superfamily 59. 12 1 60. Archeocrypticidae 1 61. 20 62. Melandryidae 38 63. 6 1 64. Rhipiphoridae 5 65. Colydiidae 196 66. 20 67. Chalcodryidae 5 68. Tenebrionidae 149 10

69. 1 70. 18 3 71. Pyrochroidae 7 72. 22 73. 17 9

74. 1 75. Scraptidae 4 Superfamily 76. Cerambycidae 180 8 77. Chrysomelidae 134 19 Superfamily CURCULIONOIDEA 78. Nemonychidae 4 79. 58 3

80. 1

81. 3 1 82. 1496 46