FNZ 43 - Carabidae (Insecta: Coleoptera) catalogue - Introduction
Larochelle, A; Larivière, M-C 2001. Carabidae (Insecta: Coleoptera): catalogue. Fauna of New Zealand 43, 285 pages.
(
ISSN 0111-5383 (print),
;
no.
43.
ISBN 0-478-09342-X (print),
).
Published 15 Jun 2001
ZooBank: http://zoobank.org/References/D7D35747-8FA5-44E4-A6AA-BD8A14204ADC
Introduction
The family Carabidae (including tiger beetles) is a highly speciose, worldwide beetle family, with an estimated total of 25,000 to 50,000 species. Carabids, or ground-beetles, are a highly adaptable group which has managed to occupy most terrestrial habitats on nearly all continents and most islands since its origin in the early Mesozoic. As a family they exhibit a relatively high degree of morphological uniformity which makes these insects especially suitable to study the ecophysiological adaptations required to cope with environmental demands. They are generally abundant in ecosystems and demonstrate a flexible set of responses to both abiotic and biotic environmental factors. Most carabids are nocturnal predators (Larochelle, 1990) living at the surface of the ground while some species live in the soil, in caves, or on the vegetation. Ground-beetles show good dispersal capability and, in many species, the ability to form stable populations following colonisation events. They are also fairly easy to sample by reliable and quantitative methods, e.g., pitfall trapping. These and other factors have made this group a favourite with biologists investigating evolutionary and ecological hypotheses throughout the world. More recently, this group has been used for applied investigations such as conservation evaluation.
The estimated number of species of better known continental faunas such as North America, Europe, and Australia, varies from about 2,500 to 3,000 species. Compared with these larger regions of the world the New Zealand fauna, currently known from five subfamilies, 20 tribes, 78 genera, and 424 species, appears relatively small, but what it lacks in size it makes up in uniqueness, e.g., 92% of known species are endemic.
The Carabidae are among the best represented groups of insects in New Zealand entomological museums and collections. Despite such high interest in this group, no modern effort has been made to catalogue the New Zealand species since Hudson's (1934) list which enumerated 556 taxa. Numerous nomenclatural changes and new taxa have been published since then and although Hudson's list is still useful, it is no longer reliable.
The present catalogue is an attempt to bring together the available literature and collection-based information on taxa recorded from New Zealand's main islands and its offshore islands. It has been written with the needs of systematists, identifiers, ecologists, and other biologists in mind, hence the sections summarizing for all species the geographic distribution, ecology, biology, dispersal power, and the citation of main references to available identification tools, taxonomic revisions, and natural history treatments. A species checklist, a full bibliography, a taxonomic index, several appendices, and species distributions maps are also provided.
All attempts have been made to report information as accurately as possible, but none are more aware than the authors of the inevitability of errors or omissions in this type of work. Therefore, the authors ask the indulgence of readers and can only hope that the usefulness of this catalogue will outweigh it shortcomings.
Brief history of carabid taxonomy in New Zealand. The first carabid beetle to have been described from New Zealand is Cicindela tuberculata Fabricius, 1775. Subsequently, between 1843 and 1923, the majority of species were described by Blanchard (1842-1843, 1853), White (1843, 1846), Laporte de Castelnau (1867-1868), Bates (1867-1878), Broun (1876-1923), and Sharp (1878-1903). This literature is quite complex to follow as about 90 species have been described twice or thrice in different papers or pre-publication reprints by the same author, e.g., Blanchard (1842-1843; 14 taxa republished in 1853), Laporte de Castelnau (1867; 23 taxa republished in 1867-1868), Bates (1874; 17 taxa republished in 1875), Broun (1882; 24 taxa republished in 1883 and 1886, and 1884; 11 taxa republished in 1886). Emberson (1993b) published an extremely useful paper establishing the publication dates of species described by Blanchard in Dumont D'Urville's Voyage au Pôle Sud.
Broun has been the most prolific carabid describer, with 382 taxa (202 still valid), but at the same time he contributed numerous nomenclatural problems still baffling workers today. His species concept was highly typological which often resulted in the description of the same species more than once under different names (e.g., 13 synonyms and 3 replacement names under Megadromus meritus (Broun)). Unfortunately, his type designations were often unclearly documented and he did not provide identification keys clarifying his view of diagnostic features. In addition, many of the Pterostichini and Platynini that he described were attributed to European genera. Examination of Broun's original material, located in the Natural History Museum (London), is essential to taxonomists studying New Zealand Carabidae. For example, among the 137 taxa described by him in the Pterostichini, only 38 are still valid after two subsequent revisions (Britton, 1940; Butcher, 1984).
Taxonomic revisions started in 1937 and the tribes treated since then are: Broscini (Britton, 1949); Bembidiini, except Tachyina (Jeannel, 1937; Lindroth, 1976 and 1980; Moore 1980b); Pterostichini (Britton, 1940; Butcher, 1984); Lebiini (Britton, 1941); Pentagonicini (Britton, 1941); and cave-dwelling carabids (Britton, 1958-1964). Based at the Natural History Museum (London), Britton contributed the greatest number of revisions. Unfortunately, even though he dealt with a good proportion of Broun's material, he did not generally designate any lectotypes. Consequently, much of his material will need to be re-examined by workers wanting to describe new taxa in the groups he previously covered.
The revisions published so far cover approximately 230 taxa which is about 50% of the currently described fauna. However, except perhaps for the genus Holcaspis (Pterostichini) which has been recently revised, most of these groups are still in need of a more modern revisionary treatment that would include the examination of all types with appropriate type designations, and a thorough study of male genitalia and other morphological features throughout the distribution range of species and of the variation of these structures within and between populations. In addition, so much new material has been collected and deposited in New Zealand collections in the last 30-40 years that numerous new taxa remain to be described in the groups already covered by previous workers as well as in other groups. We estimate that the fauna will reach around 600 species when totally described.
Most taxonomic works published until now deal only with the adult stage; the larva of only 14 species having been described: Amarotypus edwardsii Bates (Amarotypini); Bountya insularis Townsend, Brullea antarctica Laporte de Castelnau, Diglymma castigatum Broun, Mecodema alternans hudsoni Broun (Broscini); Cicindela parryi White, Cicindela tuberculata Fabricius (Cicindelini); Loxomerus brevis (Blanchard), Loxomerus nebrioides (Guérin-Méneville) (Migadopini); Megadromus vigil (White), Plocamostethus planiusculus (White), Zeopoecilus putus (Broun) (Pterostichini); Duvaliomimus mayae Britton, and Kenodactylus audouini (Guérin-Méneville) (Trechini).
Two keys to suprageneric taxa have been published: a key to carabid tribes by Britton (1940) and, more recently, a key to subfamilies and tribes by Klimaszewski & Watt (1997). A key to described genera, although an essential tool for most taxonomists and field workers, is still unavailable. Table 1 compares the subfamilies and tribes used in the present catalogue with those given by Klimaszewski & Watt (1997).
As for checklists, Hutton (1874; 1904) and Hudson (1923; 1934) are still the only lists available, now unreliable due to the numerous nomenclatural changes and new taxa proposed since.
Higher classification. The history of carabid classifications has beeen superbly presented by Ball (1979) and Ball et al. (1998b). The higher classification of carabids is very complex, with little consensus on the family's divisions, except at the tribal level. According to Ball et al. (1998b), Erwin's classification (1991) is the most detailed ever presented. His classification is very comprehensive, taking into account morphological and chemical features, behaviour of both adults and larvae, and historical zoogeography. The higher classification used in this catalogue (Table 2) follows Erwin (1991), except in the case of the Psydritae (Baehr, 1998) and the Pentagonicini (Bousquet & Larochelle, 1993). The subtribal classification, being omitted by Erwin, takes after Bousquet & Larochelle (1993), except for the Cicindelina (Freitag, 1999), the Creobiina and Broscina (Ball, 1956), the Oopterina (Liebherr & Will, 1998), and the Actenonycina (Ball et al., 1995).
More work on the systematics of Australian Carabidae is needed before the taxonomic limits and the phylogenetic relationships of the tribes Pamborini, Amarotypini, Migadopini, Broscini, Mecyclothoracini, Meonini, Tropopterini, Trechini, Zolini, and Bembidiini (Anillina), and their included subtribes, can be more satisfactorily established. Such advances will also provide the wider framework necessary to evaluate the taxonomic status of many genera originally described from New Zealand.
Geographic distribution. The New Zealand fauna is highly insular, with 46 genera (58%) and 391 species (92%) presently recorded as being endemic (Table 3).
The maps on pages 223-270 summarise the geographic distribution of Carabidae taxa occurring in New Zealand. Overall, species appear to be more widely distributed than originally thought for mostly flightless organisms; species belonging to well-revised groups have been shown to occur in several areas.
A greater number of taxa (299) occurs on the South Island, although 209 are restricted to it. Two hundred and one (201) taxa occur on the North Island, but only 111 species are restricted to it. Ninety (90) taxa are shared between the North and the South Islands.
Patterns of taxonomic diversity and the number of taxa restricted to areas of New Zealand are illustrated on Maps 4-6 (pp. 219-221). The areas so far known to contain the highest diversity (Map 4) are: NN (126 taxa), WN (121 taxa), MC (117 taxa), AK (95 taxa), BR (88 taxa), ND (85 taxa), and CO (83). Three areas of the South Island have a relatively low number of taxa: KA (44 taxa), SC (44 taxa), and MK (52 taxa). These areas have probably been undersurveyed; their number of taxa may eventually double. The areas with the greatest number of New Zealand endemics (Map 5) are found mostly on the South Island: NN (115), MC (106), WN (101), BR (85), CO (78), and DN (74).
Several carabids are restricted to a single area (Map 6). Currently, the areas with the greatest number of such taxa are: NN (19 taxa), MC (17 taxa), OL (11 taxa), ND (8 taxa), and AK (7 taxa). These numbers may change once the fauna is revised, but general trends should remain, i.e., these should be the areas of higher endemism.
Finally, the areas including the largest number of adventive taxa (Map 7, p. 222) are: AK (23 taxa), ND (18 taxa), WN (16 taxa), CL (14 taxa), and BP (14 taxa). Most introductions occur in the northern half of the North Island and in the Wellington region.
Faunal composition and affinities. Table 3 shows the total number of genera and species occurring in New Zealand. The number of adventive species is 27 or only 6% of the total fauna.
The New Zealand carabid fauna (424 species) is about 17% the size of the Australian fauna which is approximately 2,500 species according to Moore et al. (1987). It is characterised by a relatively high proportion of large-sized and darkly coloured taxa. Most species are flightless; even a few species of Cicindelini appear to have a subtantially reduced ability to fly. The largest carabid tribes in New Zealand and Australia are the Broscini and Pterostichini. The largest carabid genus in New Zealand is Mecodema (58 taxa). The following world tribes appear to have their greatest diversity in New Zealand and Australia: Amarotypini, Broscini, Mecyclothoracini, Meonini, Migadopini, Pamborini, Tropopterini, and Zolini.
Ninety-seven (97) species have been reported from New Caledonia (Heller, 1916), 15 from Norfolk Island (Moore, 1985), and 28 from Lord Howe Island (Moore, 1992). Tables 4 and 5 show the genera and species shared between these areas. Too little is known of the South American fauna to infer any relationships with the New Zealand fauna.
The New Zealand fauna shows the greatest affinity with eastern Australia (including Tasmania) in sharing 16 native genera and 4 native species. In addition Oopterus (Zolini) and Kenodactylus (Trechini) are shared with the Falkland Islands and Patagonia, but not with Australia. All Clivinini occurring in New Zealand have been introduced from Australia. Based on our preliminary examination of material in New Zealand collections and our own field work, we think that there will be additional introduced species recognised as investigations progress on the New Zealand fauna. This will probably include Australian species already introduced on Lord Howe and Norfolk Islands, e.g., Chlaenius flaviguttatus Macleay, C. ophonoides Fairmaire (Callistini); Egadroma subrobusta Schauberger, Euthenarus promptus (Erichson), Gnathaphanus melbournensis (Laporte de Castelnau), Notiobia melanaria (Dejean) (Harpalini); Eurystomis castelnaui Chaudoir (Pterostichini); Illaphanus stephensii Macleay, Paratachys transversicollis Macleay, Tachys plagiatus Putzeys (Bembidiini); and Sarothrocrepis inquinata (Erichson) (Masoreini).
Ecology, biology, and dispersal. Most New Zealand ground-beetles are hygrophilous species living at the surface of the ground, although a number of species also live deep in the soil (e.g., Anillina), in caves (see below), and also on plants and trees. Several species in the genera Oopterus, Zolus (Zolini), Molopsida (Tropopterini), "Anchomenus", Ctenognathus (Platynini), Agonocheila (Lebiini), and Amarotypus (Amarotypini) are tree-dwellers, often occurring in association with the loose bark of trees or in rotten branches.
The two native habitats harbouring the greatest number of species are forests and, to a lesser extent, tussock grasslands. In general, native carabids tend to live within the confines of native habitats, but since less than 25% of New Zealand's original forest cover remains, most species appear also to survive in modified environments. Introduced species seem to be able to invade natural habitats, but only to a slight degree. About 50 species (mostly native) dwell sucessfully in pine plantations provided these are located near or adjacent to native forests.
A number of species live almost exclusively along coastal lowlands: Cicindela (3 spp.) (Cicindelini), Brullea (1 sp.), Mecodema (3 spp.) (Broscini), Kenodactylus (1 sp.), Māoritrechus (1 sp.) (Trechini), Bembidion (2 spp.), all Zecillenus (5 spp.) (Bembidiini), Triplosarus (1 sp.) (Harpalini), and Ctenognathus (2 spp.) (Platynini). The majority of New Zealand species, however, can be found from the lowlands to higher elevations below the subalpine zone. Very few described species appear to be restricted to the subalpine and alpine zones, but many species remain to be described from such habitats, e.g., in the Migadopini and Amarotypini.
Twelve (12) species from the following genera are obligatory cave-dwellers, i.e., living exclusively in caves: Duvaliomimus, Erebotrechus, Neanops, Pholeodytes, Scototrechus (Trechini), and Syllectus (Harpalini). Eighteen (18) species are occasional cave-dwellers. They belong to the genera Taenarthrus (Migadopini), Mecodema (Broscini), Duvaliomimus (Trechini), Paratachys (Bembidiini), Holcaspis, Megadromus, Plocamostethus, Rhytisternus (Pterostichini), Dicrochile (Licinini), Lecanomerus, Syllectus (Harpalini), and Prosphodrus (Platynini). The latter two genera occur more frequently in caves than other genera, provided that rills or small brooks run through the caves.
Eleven (11) species are regarded as being synanthropic, i.e., living around human dwellings: Megadromus antarcticus (Chaudoir), a native species, and 10 adventive species in the genera Carabus (Carabini), Paratachys (Bembidiini), Laemostenus (Platynini), Perigona (Perigonini), Rhytisternus (Pterostichini), Anisodactylus, Haplanister, Harpalus, and Lecanomerus (Harpalini).
Very little is known about the life history of native Carabidae. No life-cycle study has been published so far. The breeding type of most species, i.e., the time of the year at which they reproduce, is still unknown. Population biology and locomotory activity are virtually undocumented.
Mouthpart morphology and food data suggest that most species are opportunistic polyphagous predators. Adults are generally nocturnal and active most of the year, a little less so in winter (June-August). In general, they are active from November to March, that is, the end of spring (September-November), summer (December-February), and early autumn (March-May).
Parental care, or protection of eggs and larvae by the female, has been observed in 11 species of Pterostichini: Megadromus (8 spp.), Neoferonia (1 sp.), Plocamostethus (1 sp.), and Zeopoecilus (1 sp.).
Larvae are usually fossorial and very secretive; they are rarely encountered at the surface of the ground and can be difficult to sample by hand or by pittrapping. Larvae of Bembidion, Zecillenus (Bembidiini), and Megadromus (Pterostichini), live closer to the ground surface and they are more easily found.
New Zealand carabids have number of enemies and parasites. Starlings, kiwis, spiders, and hedgehogs are major predators while magpies, asilids, kingfishers, fernbirds, stoats, feral cats, thrushes, trouts, and rats appear to be minor predators. Based on our field experience, we think that spiders could be the most important predators, especially in tussock grasslands, herbfields, and fellfields. Hedgehogs are thought to be very active predators of carabids in tussock grasslands on the South Island. As elsewhere in the world, asilid flies, or robber flies (Diptera: Asilidae), could be regular enemies of tiger beetles (Cicindelini).
A survey of specimens in the New Zealand Arthropod Collection (NZAC) reported 48 species parasitised by Laboulbeniales (Fungi: Ascomycetes). This indicates that at least 11% of total carabid fauna is subjected to such infestations. Parasitised specimens were mostly from the tribes Trechini, Harpalini, Mecyclothoracini, Tropopterini, and Zolini. Fifteen (15) species were heavily infested in the genera Duvaliomimus (Trechini), Lecanomerus (Harpalini), Mecyclothorax (Mecyclo-thoracini), Molopsida (Tropopterini), Oopterus, and Zolus (Zolini).
A similar survey revealed 137 species as being infested with mites (Acari). Therefore, 32% of all species is subjected to this type of infestation. Eighteen (18) species were heavily infested in the genera Holcaspis, Megadromus, Plocamostethus (Pterostichini), Mecodema (Broscini), and Ctenognathus (Platynini). On a world basis, and apparently also in New Zealand, mites are among the most important parasites of carabids.
In New Zealand, species infested with Laboulbeniales or mites are rain-forest dwellers, often associated with rotten wood, branches, and logs. Our survey of NZAC specimens did not reveal any teneral carabids with Laboulbeniales or mites. Before this catalogue, only one species (Kenodactylus audouini (Guérin-Méneville)) had been reported with Laboulbeniales (Rossi, 1984); no published record is available for mites. More information on Laboulbeniales and mites associated with world carabids can be obtained from Lindroth (1948), Thiele (1977), Weir & Hammond (1997), and Vigna Taglianti & Rossi (1998).
Nematodes have been found as parasites of 5 species belonging to the following genera: Mecodema (1 sp.), (Broscini), Holcaspis (2 spp.), and Megadromus (2 spp.) (Pterostichini).
The majority of New Zealand carabids are flightless, usually using running as their main mode of locomotion. In general, carabids either burrow in the litter or the soil or run away when disturbed, but a number of interesting defense mechanisms can be observed in New Zealand species. Two Cicindela species (Cicindelini) use cryptic coloration (colour pattern blending perfectly with the background) to protect themselves against enemies. Death feigning has been observed in species of Diglymma (2 spp.), Mecodema (2 spp.), Metaglymma (1 sp.) (Broscini), Megadromus (1 sp.) (Pterostichini); and possibly also Oopterus, and Zolus (Zolini) (Hudson, 1934). When disturbed, adults may emit a strong smell, e.g., Amarotypus (1 sp.) (Amarotypini), Diglymma (1 sp.), Mecodema (3 spp.), Oregus (1 sp.) (Broscini), Holcaspis (2 spp.), Megadromus (5 spp.), Plocamostethus (1 sp.), Rhytisternus (1 sp.), and Zeopoecilus (1 sp.) (Pterostichini). Species of the following genera have been observed to bite strongly when seized: Mecodema (1 sp.) (Broscini), Megadromus (4 spp.), Plocamostethus (1 sp.), and Zeopoecilus (1 sp.) (Pterostichini). When closely approached, adults of Amarotypus (Amarotypini) standing on tree trunks at night let themselves drop to the ground. Finally, Prosphodrus (Platynini) species have the peculiar habit of diving into water when disturbed.
Fifty (50) carabid taxa were listed by Molloy et al. (1994) as threatened. So far, two species (Mecodema chiltoni Broun, M. laeviceps Broun) have been the subject of a basic conservation study (Barratt, 1994a & b). In our opinion, however, too little information is available on the relative abundance and distribution of supposedly rare species; more thorough investigations, e.g., quantitative pittrap population dynamic studies and mark-recapture studies over more than one year, will need to be carried out before any meaningful formal conservation assessment can be formulated for any species. Based on the world literature and our experience in the field, ground-beetles are usually well hidden and scattered by day through a given habitat so that they usually escape the attention of the most specialised or attentive collector, thus preventing any realistic approximation of population size and distribution from hand-collecting data.