FNZ 69 - Carabidae (Insecta: Coleoptera): synopsis of species - Introduction
Larochelle, A; Larivière, M-C 2013. Carabidae (Insecta: Coleoptera): synopsis of species, Cicindelinae to Trechinae (in part). Fauna of New Zealand 69, 193 pages.
( ISSN 0111-5383 (print), ISSN 1179-7193 (online) ; no. 69. ISBN 978-0-478-34738-8 (print), ISBN 978-0-478-34739-5 (online) ). Published 7 Mar 2013
The family Carabidae (including tiger-beetles) is taxonomically diverse, with an estimated total of over 34,000 species in 1,927 genera (Lorenz, 2005). Carabids occupy most terrestrial habitats on nearly all continents. These beetles are abundant in the field and attract attention with their shape and coloration. They are mostly nocturnal and polyphagous predators (Larochelle, 1990) although some are diurnal or phytophagous. Most ground-beetles, in temperate zones at least, live at the surface of the ground, while some species live in the soil (e.g., Anillina), in caves (e.g., Trechini, Harpalini), or on the vegetation (e.g., Zolini, Lebiini). Most New Zealand carabids are flightless, which makes their dispersal capacity somewhat limited and their populations morphologically varied, sometimes even aberrant. In 2001 Larochelle & Larivière catalogued the fauna (Fauna of New Zealand 43). In 2007, they updated this inventory (Fauna of New Zealand 60) by recording 7 subfamilies, 21 tribes, 86 genera, and 461 species. The current work recognises 7 subfamilies, 20 tribes, 97 genera, and 518 species.
As a family, Carabidae exhibit a relatively high degree of morphological uniformity, making them suitable for studying the ecophysiological adaptations required to cope with environmental demands. Being sensitive to their environment, they demonstrate a flexible set of responses to both abiotic and biotic factors. Carabids are commonly used as bioindicators (Larochelle & Larivière, 2003) to assess the biodiversity of ecosystems, indicate the impact of landscape changes, evaluate environmental health, predict the effect of climate changes, classify habitats for nature protection, and characterise soil-nutrient status in forestry. They can also be used to control pest invertebrates (e.g., lepidopteran caterpillars). In the future, ground-beetles may become more commonly used in biological and integrated programs, e.g., as natural control agents of noxious invertebrates, especially soil insect pests, or control agents of weeds, especially their seeds.
This Synopsis is aimed at systematists and identifiers. Its purpose is to provide a taxonomic overview of all New Zealand species and subspecies of Carabidae, primarily in the form of an identification guide for taxonomically well-known groups or in the form of taxonomic reviews and revisions for lesser- or little-known groups. Further goals of this synopsis are to supplement the Synopsis of supraspecific taxa (Larochelle & Larivière, 2007a) and the New Zealand Catalogue (Larochelle & Larivière, 2001) by providing additions, corrections, or changes and updating the faunal checklist. The Synopsis will be published in several monographs. The present work covers the tribes Cicindelini, Pamborini, Amarotypini, Migadopini, Clivinini, Moriomorphini (formerly Mecyclothoracini, Meonini, Tropopterini), and Trechini.
This work is one more step in the authors’ goal of reaching an overall understanding of the New Zealand carabid fauna within a reasonable time frame and making relatively large amounts of information available for practical use by a wide range of end-users. The methodology is based on the concept of ‘practical taxonomy’ described by Darlington (1971), which aims to provide “a floor plan for more detailed taxonomic, ecological, zoogeographical, and evolutionary studies.”
Recent history of New Zealand carabid taxonomy
A more detailed account of the history of carabid taxonomy in New Zealand can be found in Larochelle & Larivière (2001, 2007a); recent developments are emphasised here.
The following taxonomic works have been recently published: Larochelle & Larivière, 2007b (identification guide to genera); Liebherr & Marris, 2009 (Mecyclothorax revision); Johns, 2010 (Migadopini revision); Townsend, 2010 (Trechini revision); Liebherr, 2011b (Meonochilus revision, Rossjoycea description); Liebherr et al., 2011 (Orthoglymma description); Seldon & Leschen, 2011 (Mecodema in part, revision); Townsend, 2011 (Mecodema in part; taxonomy); and Seldon et al., 2012 (Mecodema in part; revision). Finally, Will (2011) transferred the genus Cerabilia from the tribe Platynini to the tribe Loxandrini, establishing the first record of this tribe for New Zealand.
At a higher classification level, Liebherr (2011a) synonymised the tribes Amblytelini, Melisoderini, Tropopterini, Mecyclothoracini, and Meonini under the tribe Moriomorphini.
The history of carabid classification has been extensively discussed by Ball (1979), Bousquet & Larochelle (1993), Ball et al. (1998b), and Arndt et al. (2005). The classification presented by Arndt et al. (2005) takes into account the major changes brought about by recent scientific research at the subfamily level only and it is followed here. An alternative classification published by Bouchard et al. (2011) places the tribe Broscini in the subfamily Broscinae.
The tribal classification used by Larochelle & Larivière (2001, 2007a) and kept here, is based mostly on the classification proposed by Erwin (1991), which still receives general acceptance from the scientific community.
Table 1 offers a comparison between the higher classification used in the present work and in Larochelle & Larivière (2007a). Four major changes are noted. The tribe Carabini, previously recorded for New Zealand, is hereby removed; the establishment of Carabus nemoralis O.F. Müller, 1764 in New Zealand cannot be supported by collection records following its introduction from Europe in 1948. Bell & Bell (2009) and Erwin (2011) recognised the tribe Rhysodini, previously a distinct family from Carabidae, and this view is adopted here. The tribes Tropopterini, Mecyclothoracini, and Meonini were synonymised under the tribe Moriomorphini by Liebherr (2011a). The genus Cerabilia was transferred by Will (2011) from the tribe Platynini to the tribe Loxandrini.
Notes on tribes
Cicindelini. This tribe is represented by two endemic genera, Neocicindela (7 species) and Zecicindela (9 species), occurring on the North, South, and Stewart Islands. New Zealand tiger beetles are mostly diurnal and live in a range of habitats such as coastal sand beaches and dunes, riverbanks, grasslands, and roadsides. These stunning insects are recognised by the metallic colour of their body, hairy legs, large protruding eyes, strongly toothed mandibles, and the pale markings on the elytra. The most commonly encountered species are Neocicindela tuberculata (mostly North Island) and N. latecincta (South Island). Neocicindela species occur independently of water whereas Zecicindela species live along riverbanks and the seashore. A few tiger beetle species can be found in forested areas, e.g. N. parryi and N. spilleri. Sand dune species (e.g., Z. brevilunata and Z. perhispida) have cryptic coloration (body colour blending with the sandy background) and are less active during the warmest period of the day when the sand is too hot. New Zealand species fly much shorter distances than their northern hemisphere counterparts. Adults and larvae are voracious predators, feeding mostly on insects (e.g., ants). Larval development occurs in deep burrows dug into the soil. The larva positions itself at the burrow’s entrance, attached to the wall by two pairs of hooks on the dorsal surface of the abdomen, and awaits passing organisms on which it preys. Little is known about the life history and behaviour of New Zealand tiger beetles. The larvae of a number of taxa have been described.
Pamborini. This tribe is represented by a single endemic genus and species, Maoripamborus fairburni, occurring only in the northern part of the North Island. This species is nocturnal, living in forests under logs and fallen branches. This fascinating flightless insect is recognised by its elongate head, including mouthparts, adapted to feed on snails. The genus Pamborus, the only other genus in this tribe, occurs only in Australia.
Amarotypini. This tribe occurs only in New Zealand and is known from a single endemic genus and species, Amarotypus edwardsii. This group is being revised by P. M. Johns (Canterbury Museum, Christchurch), and new genera and species await description. Amarotypus edwardsii is distributed on the North, South, and Stewart Islands. This flightless beetle is easily recognised by its metallic bronze colour and its oval shape reminiscent of the Northern Hemisphere genus Amara (Zabrini). It is nocturnal, hiding during the day under the bark of live trees (e.g., Nothofagus). When disturbed, this ground-beetle drops to the ground or emits a strong smell.
Migadopini. This tribe occurs in New Zealand, Australia, the Falkland Islands, and southern South America. The New Zealand fauna comprises three endemic genera and 18 species. The genera Calathosoma and Loxomerus, with one and four species respectively, occur only on the subantarctic islands (Antipodes, Aucklands). The genus Taenarthrus, with 13 species, is distributed on the South and Stewart Islands. Species live along the edges of rills and seepages running through forests or in high altitude meadows and fellfields. They are gregarious and nocturnal, hiding during the day under embedded stones. New Zealand migadopines are flightless and predatory. The heart-shaped pronotum, suboval elytra, habitat requirements, and behaviour of Loxomerus and Taenarthrus species are reminiscent of the Northern Hemisphere Nebria (Nebriini).
Clivinini. This tribe occurs throughout the world. The New Zealand fauna is known from four species belonging to the genus Clivina introduced from Australia and occurring on the North and South Islands. The highly diverse Australasian fauna of Clivinini is being revised by M. Baehr (Zoologische Staatssammlung, München, Germany). Results from his research may affect the composition of the New Zealand fauna. Clivina species are recognisable by their pedunculate body, protibiae with a finger-like protrusion on outer side, and strongly developed mandibles. They are gregarious and nocturnal, living in wet or moist areas, hiding in burrows during the day, and flying readily to lights at night.
Rhysodini. This tribe, long regarded as a family distinct from Carabidae, occurs throughout the world. The New Zealand fauna has been revised (Bell & Bell 1978, 1979, 1982, 1985; Emberson, 1995). Four genera and six species are known. All species are endemic but the genus Kaveinga is Australasian and the genus Rhyzodiastes is known from Australasia and South America. Rhysodine species are darkly coloured, have cylindrical pedunculate bodies, and moniliform (bead-like) antennae. They also possess unique mouthparts with retractile palpi and mandibles forming a cover protective to other mouthparts rather than being used for feeding (Bell, 1994). Of the six New Zealand species, five are restricted to the northern part of the North Island; Kupeus arcuatus extends to northern areas of the South Island. Rhysodines can be found in fallen logs, erect dead trees, stumps, woody roots, and under loose bark; they are often found in relatively firm wood requiring splitting with an axe to collect them (Bell & Bell, 1995). They do not dig burrows but push themselves through the wood layers using the margin of their elongate mentum as a cutting blade (Bell, 1998). They have been observed feeding on slime molds (Myxomycetes) (Bell & Bell, 1995).
Moriomorphini. This tribe occurs in the Australian Region, on Pacific Islands, and in southern South America. The New Zealand fauna is composed of seven genera and 48 species previously assigned to the tribes Mecyclothoracini, Meonini, and Tropopterini until they were synonymised with the tribe Moriomorphini by Liebherr (2011a). All taxa are endemic except for the adventive Australian species Mecyclothorax ambiguus. Moriomorphine species are flightless (except Mecyclothorax ambiguus and M. rotundicollis), nocturnal, and probably predatory (as suggested by mouthpart morphology). Very little is known about the life history and behaviour of New Zealand moriomorphines. Their representation in collections is rather poor; many species are known from fewer than ten populations, suggesting that new taxa may be discovered with additional field surveys and specialised sampling methods (e.g., leaf-litter sifting and soil-washing).
The four species of Mecyclothorax occur on the North and South Islands, and on some off-shore islands (Kermadecs, Three Kings, Chathams). They inhabit forests, fields, sand dunes, and the vicinity of streams. These beetles are often found during the day in leaf-litter and under fallen branches. Mecyclothorax ambiguus is often seen in large numbers at the base of Lupinus plants growing on sand dunes.
The genus Meonochilus, with six species, is restricted to the North Island. These beetles are forest-dwelling and active at night on mossy logs or trees. They hide during the day under logs, fallen branches, stones, and in leaf-litter.
The genus Rossjoycea is known from a single species occurring in one locality in southern Westland, South Island. Rossjoycea glacialis is the largest New Zealand moriomorphine species (9.2–10.3 mm in length). It has been collected in shrubby-grassy rocky areas above the Franz Josef glacier. The bilobed pro- and mesotarsi suggest plant-climbing abilities.
Seven species of Selenochilus and six species of Molopsida are found on the North Island and in northern areas of the South Island. Selenochilus species superficially resemble Northern Hemisphere representatives of Phonias, a subgenus of Pterostichus (Pterostichini). Selenochilus species are unique among New Zealand moriomorphines in having verticillate setae medially on the second antennal segment. Their enlarged and heavily ciliate maxillae suggest specialised feeding, possibly on millipedes. They live in forests where they can be found during the day mostly under logs and stones. Selenochilus hutchisonae new species has been collected in large numbers by pitfall trapping. Molopsida species are also forest-dwellers. They are mostly associated with wood, hiding during the day in and under fallen rotten branches and logs. Some species occasionally climb on trees. Most taxa have strongly convex and ovate elytra, giving them a “hump-backed” appearance.
Tarastethus (six species), previously a junior synonym of Molopsida, is restricted to the South Island. Members of this genus are active at night on trees and logs. They are primarily associated with wood, hiding during the day under the loose bark of fallen tree-trunks as well as in and under logs. The bilobed pro- and mesotarsi of Tarastethus may be associated with their climbing abilities.
Trichopsida new genus, is known from 18 species, several of which had been previously assigned to Molopsida. Trichopsida occurs in southern areas of the North Island and on the South Island. Species are found in forests where they live deep in the soil, in the thick leaf-litter, or under embedded stones. This genus is unique among New Zealand moriomorphines in having palpi with setose terminal segments. Many species have strongly reduced eyes, inflated tempora, pale and flattened body, and long pubescence, suggesting subterranean habits similar to those of Anillina (Bembidiini) that live deep in soil fissures and in thick leaf-litter. Some species of Trichopsida can be collected only by using soil-washing techniques.
Trechini. This tribe is represented by 11 genera and 34 flightless species distributed from the North Island to the subantarctic islands. All taxa are endemic to New Zealand, except Kenodactylus audouini which also occurs on the Falkland Islands, South Georgia Island, and in Patagonia. A high proportion of New Zealand trechines are cave-dwellers; many species are riparian (e.g., most Duvaliomimus species); some species live in the vicinity of seashores under stones and among gravel (e.g., Kenodactylus, Maoritrechus, Oarotrechus); one species (Kiwitrechus karenscottae) is found in the humus and leaf-litter of Nothofagus rainforests. Cave-dwelling species are usually pale in colour, flat-bodied, without eyes, and with long antennae and legs. Trechine species are probably predacious (as suggested by mouthpart morphology). The most diverse genus is Duvaliomimus with 13 species and two subspecies. Members of this genus are gregarious and nocturnal, hiding during the day under stones and among gravel.