MÄURI ORÄ, 1980, 8: 55-67 55

GROÜND (CARABIDAE) OF ARTHURS PASS

NATIONAL PARK

P.M. JOHNS

Department of Zoology, University of Canterbury, Christchurch, New Zealand

ABSTRACT

Fifty species of Carabidae are recorded from Arthurs Pass National Park. One synonymy is established and several species and one genus have been found to be undescribed, but these are left to await formal reviews of their groups. The species present have been placed in several groupings or faunules, these being related to the general environment based mainly on * rainfall and Vegetation cover. The main populations of the species of these faunules lie mainly to the north and the small populations of some species could be considered as relicts.

INTRODUCTION

Arthurs Pass National Park (Fig. 1) comprises about 100,000 ha of mountains and glacial Valleys. Although the Valleys may have flat floors at about 700m a.s.l. in the east and 300m in the west, the topography is rough with 21 peaks rising above 1800m and ten over 2100m. The Main Divide is within the western portion of the Park. Most of the eastern slopes and flats are covered in an almost uniform mountain beech forest (Nothofagus solandri var. cliffortioides) with minor pockets of N. fusca and N menziesii. In the high rainfall western portion a diverse podocarp forest dominates at lower levels and this passes into rata/mountain totara forest above. The upper margin of the forest is at about 1200m and the area of subalpine to alpine scrub, tussock and rock zones is about equal to that of the forest below. Arthurs Pass National Park has long been a collecting area for entomologists. Its diverse habitats, from rainforest on the west, to the central highland swards, screes and crags, thence to the dry forests, tussocks and river beds of the east, have been sources of a wide variety of . A preliminary checklist records 523 species of arthropods from the area (Johns unpub.) and of these 137 species have the Park as their 56 MÄURI ORÄ, 1980, Vol. 8

Fig. 1. Location of Arthurs Pass National Park in the high country of central South Island, New Zealand. type locality. The percentage, 26%, is the highest known in New Zealand and it could be rivalled perhaps only by the unknown percentages of types from Tongariro National Park or the Mt Arthur region of Nelson. This percentage could be raised if the species of the nearby Cass, Craigieburn and Porters Pass areas are added. A list for the nearby Cass Biological Station area (Anon 1977) was provided in part by the author and the distributions and biologies of some of the species have been reported (Johns 1977). Some notes are also provided in the "Natural History of Canterbury" (Johns 1969) and this book, along with the Arthurs Pass National Park Handbook (Burrows 1974), review the general geology, geomorphology, soils and Vegetation of the area. The biologies of some species of carabids complement the biologies of the millipedes of the area, especially those of the lowland Canterbury fauna which extends into the intermontane Valleys (Johns 1970, 1979). JOHNS - ARTHURS PASS CARABIDAE

CHECKLIST OF ARTHURS PASS NATIONAL PARK CARABIDAE (Subfamilies as in Britton 1970)

Migadopinae Amarotypus edwardsi Bates, 1872 Loxomerus capito Jeannel., 1938 New genus and species

Broscinae T rugiceps rugiceps Sharp, 1886 Mecodema metallicum Sharp, 1886 Mecodema costellum lewisi Broun, 1908 Mecodema allani Fairburn, 1945 Mecodema new species Mecodema fulgidum Broun, 1881 Diglymma clivinoides (Castelnau, 1867) Metaglymma monilifer Bates, 1867

Zolinae Zolus helmsi Sharp, 1886 Oopterus latipennis Broun, 1903 Oopterus parvulus Broun, 1903 Oopterus laeviventris (Sharp, 1886) Oopterus lewisi (Broun, 1912) Oopterus new species

Bembidiinae Bembidi on mao rinum Bates, 1867 Bembidi on deh iscens Broun, 1893 Bembidi on cha rile Bates, 1867 Bembidi on tai ruaense Bates, 1878 Bembidi on cha lceipes Bates, 1878 Bembidi on ach onoderum Bates, 1878 Bembidi on hok itikense Bates, 1878 Nesambl yops o reobius (Broun, 1893) Psydrinae T Solenochilus syntheticus (Sharp, 1886) T Molopsida debilis (Sharp, 1886) Molopsida puncticollis (Sharp, 1883) Mecyclothorax rotundicollis White, 1846

Pterostichinae lobipes (Bates, 1878) (Chaudoir, 1865) Megadromus enysi (Broun, 1882) Megadromus new species Neoferonia ardua (Broun, 1893) Holcaspis hudsoni Britton, 1940

Agoninae Agonum helmsi Sharp, 1886 Agonum integratus Broun, 1908 Notagonum submetallicum (White, 1846) Notagonum feredayi (Bates, 1878) Zabronothus Oblongus Broun, 1910 Colpodes haasti(Broun, 1886)

Licininae Dichrochile flavipes Broun, 1917 Dichrochile thoracica Broun, 1908

Harpalinae Syllectus anomalus Bates, 1878 Hypharpax abstrusus Bates, 1878 58 MAURI ORA, 1980, Vol. 8

Lebiinae dieffenbachi White, 1843 Demetrida nasuta White, 1846 Pentagonicinae versicolör Bates, 187 8 Scopodes laevigatus Bates, 1878 Odacanthinae Actenonyx bembidioides White, 1846 T indicates that the species has A.P.N.P. as its Type Locality.

HABITAT AND DISTRIBUTION

The carabid fauna of New Zealand has not been completely reviewed and few keys are available. Britton (1940, 1970) gives keys to the subfamilies and has reviewed the (1940), (1949) and the and Pentagonicinae (including Odacanthinae) (1941). The Migadopinae has been reviewed by Jeannel (1938) but more than half the species of this subfamily in New Zealand are known to be undescribed and this work covers only some of the common ones. A review of the Zolinae (syn. Merizodiinae (Johns 1974)) and a review of the genus Megadromus are in preparation (Johns in prep.). The Agoninae, Psydrinae (syn. Nomiinae) and Licininae are all in need of basic revisions although most of the Arthurs Pass species are easily identified through their original descriptions.

AMAROTYPUS EDWARDS! BATES, 1872. This is a remarkable member of the New Zealand carabid fauna and is one which is confined to the trunks of trees, predominantly in Nothofagus forests. Although the genus is monospecific at present there is a further species in the alpine zone of north Westland and western Nelson. It is widespread in the South Island and moderately common in most of the Arthurs Pass forests. There are no confirmed records of it being taken away from living trees. On suitable nights it may be seen wandering on trunks, among the mosses and lichens. Common associates are Artystona spp. (Tenebrionidae), Rhygmodus spp. (Byrrhidae), Celatoblatta spp. (Blattodea) and rhaphidophorid wetas (Orthoptera: Rhaphidophoridae). Larvae of Artystona spp, young cockroaches and wetas, and collembola may well be the prime food of this carabid.

It is virtually confined to Nothofagus forests on the main islands but it is known from kamahi-kohekohe forest on Inner Chetwode Island and kamahi-podocarp forest on Stewart Island. Nothofagus is absent from both these islands.

LOXOMERUS CAPITO JEANNEL, 1938. The record of this species from Arthurs Pass could well be its northernmost for it is a rare, generally subalpine, of western Otago, south Westland and Southland. At Arthurs Pass it is known only from two specimens taken from the edge of the Otira Glacier stream at 1220m, its immediate habitat being the mosses beside the stream. A similar habitat has been noted for it in Westland National Park (Townsend pers. comm.), but at Coronet Peak and further south it has been JOHNS - ARTHURS PASS CARABIDAE 59

taken near rocky streams in Nothofagus forest and even near sea level in the far south. There is no apparent morphological Variation between these populations. Its close relative, Loxomerus brevis of the subantarctic Auckland Islands, is also found in a similar ränge of habitats (Johns 1974). * MIGADOPINAE NEW GENUS AND NEW SPECIES. A very small which is known only from Temple Col (1567m) and nearby. It was found under fine stones beside large rocks or crags and where there was very little Vegetation nearby. A few Celmisia plants were present. Other undescribed species of the same genus are known from similar high alpine screes of South Canterbury, and this species may be also represented by two specimens from Mt Miromiro near Hanmer Springs.

MECODEMA RUGICEPS RUGICEPS SHARP, 1886. This species is perhaps the characteristic carabid of the wet, mountain forests of the South Island. However, in Arthurs Pass National Park it is absent from most of the higher forest and subalpine scrub and is rare in the drier, eastern portions. It is more common in the relict Stands of Nothofagus fusca - N. menziesii forest. Its füll distribution is given by Townsend (1965) . The original specimens were taken near "Bealey" in 1884. MECODEMA METALLICUM SHARP, 1886. This is usually found in the lowland podocarp-Metrosideros forests of Westland and the one record of the species from the very western part of the Park is consistent with this. -* MECODEMA COSTELLUM LEWISI BROUN, 1908. Nowhere is this species common (Townsend 1965). Within the Park it is confined to a few small terraces which are probably relict portions of the shores of Glacial Lake Speight, a large lake which once filled * the Waimakariri Valley between the present Hawdon and Poulter Rivers. It is a burrower and its burrow usually opens near the edge of a log or into the space under a log. This subspecies, formerly given füll species rank, is virtually confined to the Park (Townsend 1965).

MECODEMA ALLANI FAIRBURN, 1945. This is one of the rarities of the Park and its populations are small and isolated. It was first described from Pylon Gully, near Cass and within the Park it has been found only in the Andrews Valley, Hallelujah Fiat and near the Trust Hut in the Poulter Valley. Within and to the north of the Park (Hurunui Valley and Lewis Pass) this species is generally associated with the Nothofagus fusca - N. menziesii forest but it is not restricted to this type of Vegetation. Its southern populations (recorded by only three specimens at Pylon Gully and in the Wilberforce Valley) are certainly within pure Stands of Nothofagus solandri v. cliffortioides.

MECODEMA NEW SPECIES. This species is only slightly smaller than Mecodema costellum but it has a very different habitat, being confined to the relict Stands of Nothofagus fusca - N. menziesii m in the eastern portion of the Park. It lives within large, rotten logs. 60 MÄURI ORA, 1980, Vol. 8

MECODEMA FULGIDUM BROUN, 1881. This carabid is characteristic of the dry Nothofagus forests and scrub of the eastern portions of the South Island north of the Rangitata River. It is also present in northwest Nelson and Marlborough.

DIGLYMMA CLIVINOIDES (CASTELNAU, 1867). Another beetle commonly found in the wetter Nothofagus forests of the South Island. Present throughout the Park west of the approximate 1500mm isohyet.

METAGLYMMA MONILIFER BATES, 1867. Although characteristic of the dry scrublands of Canterbury, within the Park this species is confined to the edges of the Nothofagus forest east of the Hawdon Valley. It is, however, rare even there. It is often associated with Megadromus antarcticus.

ZOLUS HELMSI SHARP, 1886. Arthurs Pass National Park may be close to the southern limit of this northwest Nelson - Westland species and this is reflected in its abundance in the sheltered warmer parts of the Park. High populations have been seen not only in the Nothofagus fusca - N. menziesii Stands but also in the Hawdon Valley, Klondyke Corner and Kellys Creek. It is seldom seen at higher levels, even at the township (750m). Its eastern limit approximates the 1500mm isohyet.

OOPTERUS SPECIES. The type of Tarastethus lewisi Broun, 1917 (held in the British Museum, Natural History) clearly shows that this species is a zoline of the genus Oopterus (New Combination). The four species, O. lewisi, O. laeviventr is, o. parvulus and O. latipennis are all similar in size and occupy very similar habitats and all have been taken from at least one locality, Kellys Creek. There these species live under logs and stones in the scrub on the flats by the creek, with O. latipennis being the most common. This species also extends farther eastwards and reaches to higher levels on the slopes, judging from the few records available from within or near the Park. Nothing further can be said about the other species for they are known by only a few specimens from the Westport-Greymouth-Reefton area immediately northwest of the Park.

OOPTERUS NEW SPECIES. Known only from one specimen taken at 1800m on Phipps Col in a fine, fractured pebble environment at the top of a scree. A few Celmisia plants were nearby and the specimens of the new genus of migadopine were also collected nearby.

NESAMBLYOPS OREOBIUS (BROUN, 1893). This minute beetle was collected in the Otira Gorge and the Single specimen is still the only one known from the area. That specimen was described as the species Tachys coriaceus Broun, 1908 and of course, Otira Gorge is its type locality. However, Moore (1980) now considers this name to be a synonym and the distribution of the species is throughout the North Island, Nelson and northern Westland, with the Otira record being the southernmost. JOHNS - ARTHURS PASS CARABIDAE 61

SOLENOCHILUS SYNTHETICUS (SHARP, 1886). Without examining the types, Britton (1940) synonymised Solenochilus piceus (Blanchard) and Sympiestus syntheticus Sharp. This was accepted by Johns (in: Anon. 1977) in recording it from the Cass area. There are howeverf small but consistent differences between the two and for the present they are regarded as separate species. Solenochilus piceus would thus be endemic to Banks Peninsula and s. syntheticus would be the name for the species along the main alpine chain. The type specimens of s. syntheticus were taken from "Bealey", presumably near the coach stop in 1884 and the species has since been collected in the vicinity and at Klondyke Corner. The types of another species, s. fallax, were taken at Castle Hill Station, some 30km east and this is probably a synonym of the above. The species as a whole seems to be confined to the drier, intermontane forests of Canterbury, however there are very few records of it.

MOLOPSIDA DEBILIS (SHARP, 1886). This is another rare beetle. It was originally collected, along with Solenochilus syntheticus at "Bealey" and has since been found at Woolshed Hill and just outside the Park at Cass and in the Wilberforce Valley. Nothing is known of its habits but the species has greatly reduced eyes in comparison with other members of the genus, thus it probably lives within the stony soils of the steepland Nothofagus forest.

MOLOPSIDA PUNCTICOLLIS (SHARP, 1883). This widespread species inhabits the high rainfall forests of northern Westland and the alpine chain between at least the Rakaia-Hokitika watershed and north to the vicinity of Lewis Pass. It is common in the wet forest of Arthurs Pass National Park, especially to the west. Its eastern boundary probably approximates the 1500mm isohyet. A small species, and as it generally occurs in crevices under logs, it is easily overlooked.

MECYCLOTHORAX ROTUNDICOLLIS WHITE, 1846. Relatively widespread in New Zealand, this beetle favours damp habitats in a wide ränge of lowland situations. It may be found under logs in forest or scrub, and under cover in grassland or riverbed.

MEGADROMUS LOBIPES (BATES, 1878). A widespread but uncommon carabid mainly of wet Nothofagus forest in the northwestern half of the South Island. In the Park it is rarely found outside the relict Nothofagus fusca - N. menziesii forests where it dwells within the large fallen logs.

MEGADROMUS ANTARCTICUS (CHAÜDOIR, 1865). Although widespread in Canterbury, within the Park this beetle is confined to relatively dry, deep loess-derived or silty soils on terraces. It has been found only on the forest edge east of the Hawdon mouth, and for Short distances, up the Poulter and Esk Valleys. At these sites it burrows under logs to 0.5m depth and is usually stationed in a short tunnel leading to the burrow or to the outside under the log. It is often associated with Metaglymma monilifer, another dry scrubland species. MEGADROMUS ENYSI (BROUN, 1882) AND MEGADROMUS NEW SPECIES. These two species are very similar in size, habits and habitat. Their distributions within the relatively dry, eastern mountain beech forests are separated by the Waimakariri River, Megadromus 62 MÄURI 0RA, 1980, Vol. 8

enysi to the south and Megadromus n. sp. to the north. Both make Short burrows (less than 10cm) under logs and are often found within cracks in logs.

NEOFERONIÄ ARDUA (BROUN, 1893). This northwest Nelson-Westland species reaches its southern limit at Arthurs Pass National Park. It has been taken in the Deception River and Kellys Creek Valleys. It is typically a beetle of high rainfall, Nothofagus or mixed Nothofagus-Podocarp forests.

HOLCASPIS HUDSONI BRITTON, 1940. This species is distributed from Kaikoura south to the very eastern portion of the Park where it is rare. It is a dry forest species and its close relative, H. suteri is found on Banks Peninsula.

ZABRONOTHUS OBLONGUS BROUN, 1910. This carabid is another rarity of the Park. It was first described from Broken River and has since been found in various areas of inland North Canterbury. Its general habitat appears to be forest {N. solandri v. cliffortioides) or scrub {Leptospermum-Pittosporum) and is found there under stones or logs. Only one specimen has been taken in the Park; from the Poulter Valley. The eyes of this species are much reduced and it is thought to be a burrower in fine loess soils. A close relative endemic to Banks Peninsula lives in a similar habitat.

AGONUM HELMSi SHARP, 1886. An uncommon species confined to the wet forests and scrublands of Westland. Within the Park it has been found only in the very western part near Kellys Creek.

AGONUM INTEGRATUS BROUN, 1908. A widespread beetle in Canterbury foothills and mountain Valleys north of the Rakaia River System. It is most common in the mountain beech forest edge.

COLPODES HAASTI (BROUN, 1886). This is rather an uncommon yet widespread species which frequents many types of relatively open tussock, scrub or forest edge habitats. It has been taken only once in the area, on the very edge of the Park near the Mt. White Bridge at Cass. It has been taken in most of the intermontane Valleys of Canterbury.

DICHROCHILE FLAVIPES BROUN, 1917. Another of the species distributed through the Nothofagus forests of the northwestern portion of the South Island and, like Mecodema allani and Megadromus lobipes, it reaches its southern limit at the Pass. It too, has been taken commonly only in the relict Stands of Nothofagus fusca - N. menziesii Stands in the eastern portions of the Park, but it is also known from the scrub along Kellys Creek.

DICHROCHILE THORACICA BROUN, 1908. First described from the Craigieburn Range this beetle is widespread in western Canterbury and Marlborough, ranging through many habitats from stony beaches through lowland scrub, riverbeds, beech forest to alpine screes. It is also known from exotic forests at Hanmer Springs. Although it prefers damp situations to this dry northeastern part of the South Island, it does not extend far into the wetter parts of the Park beyond approximately 1250mm isohyet. JOHNS - ARTHURS PASS CARABIDAE 63

SYLLECTUS ANOMALUS BATES, 1878. A widespread but uncommon species of wet areas, S. anomalus is known from alpine streamsides to lowland floodplains especially on the West Coast. One specimen was collected at Kellys Creek.

HYPHARPAX ABSTRUSUS BATES, 1878. A common beetle which is found in many types of scrubland habitats, even when much modified for agriculture or horticulture. It is probably common to both Australia and New Zealand. In the Park it is seen only on the grassed or scrub areas of river flats.

SCOPODES VERSICOLOR BATES, 1878. One of the widespread riverine beetles of Canterbury, s. versicolor is especially abundant on the mat plant areas of the upper reaches of all Canterbury river Valleys. It scuttles over these mat plants feeding on minute and searching for mates. It is often bright coppery or greenish in colouration. Recorded also by Britton (1941) .

SCOPODES LAEVIGATUS BATES, 1878. Individuais of this species are smaller in size and are black, in contrast with s. versicolor. It does however, have much the same habitat but extends to higher levels and is often found close to glacier-fed streams or ice and snow patches.

ACTENONYX BEMBIDIOIDES WHITE, 1846. This is an abundant insect of the open river beds of the South Island. It preys on small flies and its larva is also a predator of the larvae of those flies which live in the fine muds and organic matter which accumulate around the larger stones of the river flats. The adult is a very fast runner.

DISCUSSION

Some 338 species of carabid ground beetles are recorded from New Zealand although more are yet to be described. Thus Arthurs Pass National Park with its 50 species has about 15% of the total New Zealand fauna. This richness reflects the variety of habitats within the Park ranging from high rainfall forest, through alpine scrub scree and rock to the lowland dry forest, scrubland and bare, stony river beds. Its richness is also due to the presence of relict patches of Nothofagus fusca - N. menziesii forest amongst the N. solandri v. cliffortioides forest. The fauna of the relict patches is one clearly associated with northwest Nelson, and within the Park these lie east of the Main Divide approximately between the 1500 and 2500mm isohyets. From both the west and the east the lowland species of Westland and Canterbury respectively, are distributed up the montane Valleys tili they reach their limits controlled by various environmental and biological factors which on general field observations seem related to precipitation. Some species reach further than others but it is those of the eastern, dry Canterbury faunule which have very distinct boundaries. That fauna is exemplified in the Carabidae, by Metaglymma monilifer, Megadromus antarcticus and Agonum integratum and by various 64 MÄURI ORA, 1980, Vol. 8 millipedes (Johns 1979). The Park is also the southern terminus of a complex set of species representing two distinct faunules extending from the very north of the South Island. The first of these faunules, the northwestern faunule, is associated with the relict patches of Nothofagus fusca - N. menziesii forest lying to the east of the Main Divide in the region of the Park but becomes far more extensive in the Lewis Pass and Reefton-Inangahua areas and of course it is very extensive in northwest Nelson. The species involved are listed in Table 1. The second northern faunule is that associated with the dry mountain beech forests and scrublands of inland North Canterbury and Marlborough. It has many species in common with the lowland Canterbury faunule too and perhaps the two should be regarded as subsets of the one faunule. The species represented are listed in Table 1, together with other species which do not quite reach the Park. On the very southern edge of the Park and including the northern Valleys of the Rakaia and Wilberforce Rivers there are extensive areas of Podocarpus halli (mountain totara) scrub and Nothofagus solandri forest. This region contains the rare species Molopsida debilis and Molopsida diversa, and it is suspected that these are relicts of a past, colder climate and environment, surviving in a refugium between the termini of the extensive Pleistocene Valley glaciers of the Rakaia and Waimakariri Valley Systems. Both have reduced eyes and their habits suggest that they are stony-soil dwellers, rather than litter inhabitants like most of their congeners. This area also has the endemic soil-dwelling millipede Dityloura dealbata (Johns 1970) . Thus Arthurs Pass National Park is a meeting place of at least three separate faunules (Fig. 2). I believe that nowhere eise in New Zealand can such a clear demarkation of faunules be examined. From a scientific point of view such habitat differentiation in such a small area is extremely interesting and the conservation of these habitats, especially the relict and marginal ones, is essential. Unfortunately these habitats are in areas much used by Park visitors and the siting of accommodation huts and other amenities at the Hawdon Valley entrance and at Andrews Stream may lead to a gradual deterioration of the forest margins in the vicinity with the possible consequence of extinction of the relict and marginal populations there. The upper Poulter Valley no longer has vehicle access and the distance deters all but the keen trampers. Studies on the species present in these marginal areas need not however, be concentrated on populations within the Park for all but one are found elsewhere, some abundantly so. The Cass Biological Station area is eminently suitable for a study of the Canterbury lowland faunule, whilst the Hurunui Valley and Hanmer Springs areas are more suitable for those whose major distribution areas lie to the northeast. Species of the relict Nothofagus fusca - N. menziesii forest are more common in the upper Hurunui Valley and Lewis Pass areas where this type of forest is far better developed. However, the alpine and mountain beech biotopes of the Park are extensive and, coupled with their easy access, are ideal areas for scientific studies. TABLE 1. SPECIES COMPOSITION OF CARABID FAUNULES OF ARTHURS PASS NATIONAL PARK. ARROWS INDICATE LIMITED INCURSION INTO ADJACENT FAUNULE. *, SPECIES RECORDED CLOSE TO THE PARK.

Western faunule Northwestern faunule Northeastern faunule Eastern faunule

Vegetation lowland podocarp Nothofagus fusca/ Nothofagus solandri N. solandri v. cliffortioides/ N. menziesii v. cliffortioides open scrub

Restricted Mecodema metallicum Mecodema allani • Mecodema fulgidum- species Mecodema costellum lewisi- Mecodema new species Metaglymma monilifer Zolus helmsi Oopterus parvulus Oopterus laeviventris Oopterus lewisi 1 Oopterus latipennis - Co Duvaliomimus walkeri* new genus* Nesamblyops oreobius Molopsida sulcicollis* — Molopsida puncticollis Solenochilus syntheticus Neoferonia ardua Agonum integratum I Agonum helmsi a Megadromus lobipes Megadromus n. species Megadromus enysi I Megadromus antarcticus Co Holcaspis hudsoni Zabronothus Oblongus - Zabronothus major* Dichrochile flavipes Dichrochile thoracica -

Widespread Diglymma clivinoides S.I. species Mecodema rugiceps Amarotypus edwardsi 66 MÄURI ORA, 1980, Vol. 8

LIMIT OF NORTHWESTFRN FAUNULE

WAIMAKARIRI RIVER

C RA IGIE BU RN ^llll FOREST PARK

Fig. 2. Distribution of carabid beetle faunules in the vicinity of Arthurs Pass National Park. Light stipple, eastern and western limits of western and eastern-northeastern faunules respectively. Heavy stipple, extent of northwestern faunule in this region (schematic). JOHNS - ARTHURS PASS CARABIDAE 67

ACKNOWLEDGEMENTS

I thank the Chief Rangers and the Arthurs Pass National Park Board for permission to collect within the Park on many occasions. I am especially grateful to Alan Perrett for his help during his period as a Ranger there.

LITERATURE CITED

AN0NYM0US 1977. Checklist of the Fauna of Cass. Pp. 367-399. In: Burrows, CJ. (Ed.). BRITTON, E.B. 1940. The Carabidae (Coleoptera) of New Zealand. I. The Pterostichini. Transactions of the Royal Society of New Zealand 69(4): 473-508. BRITTON, E.B. 1941. The Carabidae (Coleoptera) of New Zealand. II. Tribes and . Proceedings of the Royal Entomological Society of London (B) 10(10): 185-196. BRITTON, E.B. 1949. The Carabidae (Coleoptera) of New Zealand. III. . Transactions of the Royal Society of New Zealand 77(4): 533-581. BRITTON, E.B. 1970. Coleoptera. Pp. 495-635. In: Insects of Australia. Division of Entomology, Commonwealth Scientific and Industrial Organisation. Melbourne University Press. Melbourne. 1029 pp. BURROWS, CJ. 1974 (Ed.). Handbook to the Arthurs Pass National Park (3rd Ed. revised). Arthurs Pass National Park Board, Christchurch. 104 pp. BURROWS, CJ. 1977 (Ed.). Cass: History and Science in the Cass District, Canterbury, New Zealand. Botany Department, University of Canterbury, Christchurch. 418 pp. JEANNEL, R. 1938. Les Migadopides (Coleoptera; ). Revue Francaise d'Entomologie 5: 1-55. JOHNS, P.M. 1969. The Mountain Invertebrate Fauna. Pp. 391-39. In: Knox, G.A. (Ed.) The Natural History of Canterbury. A.H. & A.W. Reed, Wellington. 620 pp. JOHNS, P.M. 1970. New Genera of New Zealand Dalodesmidae. Transactions of the Royal Society of New Zealand, Biological Sciences 12(20): 217-237. JOHNS, P.M. 1974. Arthropoda of the Subantarctic Islands of New Zealand 1. #(Coleoptera: Carabidae). Southern New Zealand, Patagonian and Falkland Islands insular Carabidae. Journal of the Royal Society of New Zealand 4(3): 283-302. JOHNS, P.M. 1977. The Biology of the Terrestrial Fauna. Pp. 311-328. In: Burrows, CJ. (Ed.). JOHNS, P.M. 1979. Speciation in New Zealand Diplopoda. Pp. 49-57. In: Marina Camatini (Ed.) Myriapod Biology. Academic Press, London. 456 pp. LINDROTH, C.H. 1976. Genus Bembidion Latrielle in New Zealand; a revision. New Zealand Journal of Zoology 3(2): 161-198. MOORE, B.P. 1980. A Synopsis of the New Zealand Anillini (Coleoptera; Carabidae; Bembidiinae) with descriptions of new genera and species. New Zealand Journal of Zoology 7 (3): 399-406. T0WNSEND, J.I. 1965. Notes on the genus Mecodema with descriptions of new species from the South Island of New Zealand. New Zealand Journal of Science 8(3): 301-318.