FNZ 53 - Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae) - Introduction
Larochelle, A; Larivière, M-C 2005. Harpalini (Insecta: Coleoptera: Carabidae: Harpalinae). Fauna of New Zealand 53, 160 pages.
(
ISSN 0111-5383 (print),
;
no.
53.
ISBN 0-478-09369-1 (print),
).
Published 04 Jul 2005
ZooBank: http://zoobank.org/References/5A8BF6BE-EA73-4476-8C81-E4F00709AC9B
Introduction
The tribe Harpalini belongs to the subfamily Harpalinae (Coleoptera: Carabidae) which contains over 19 000 taxa. Molecular sequence data indicate that Harpalinae radiated in the Cretaceous Period (Ober 2002).
The Harpalini form a diverse group, including over 240 genera and subgenera, and approximately 2 000 species distributed in all biogeographic regions of the world. The present faunal review records 20 genera and 57 species for New Zealand. This should constitute the near totality of the fauna.
Compared with New Zealand, the Australian Harpalini are more diverse with over 160 species distributed in about 20 genera (Moore et al. 1987), but the fauna remains largely unrevised.
The present work offers a concise faunal taxonomic revision of the New Zealand Harpalini, based on the study of adults contained in local and overseas collections. It represents a first modern attempt to bring together the scattered information dealing with the group.
The goals of this revision are straightforward: to provide an inventory of New Zealand taxa, a concise treatment of their taxonomy, identification keys to genera and species, and a summary of available information on species distribution, ecology, biology, and dispersal power.
It is one step in the authors’ overall goal of attaining an overall understanding of the carabid fauna within a reasonable time frame, and to make relatively large amounts of information available for practical use by a wide range of end-users. The methodology involves less gamma taxonomy but more intensive field work, and it 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.”
It is hoped that this kind of faunal taxonomy will provide solid foundations for studies of other types, much in the same way as the work done by Lindroth between 1961 and 1969 for Canada and Alaska, and Darlington between 1962 and 1968 for New Guinea.
In addition to paper-based publications the authors publish the New Zealand Carabidae website (http://www.landcareresearch.co.nz/) which maintains up-to-date information on New Zealand carabids, including digital images, recent literature, as well as additions and corrections to previous publications.
Taxonomic history
There has been little work done on the faunistics of the New Zealand Harpalini since the earliest descriptions of Hypharpax antarcticus and Triplosarus novaezelandiae by Laporte de Castelnau in 1867. No identification keys or taxonomic overview (except for the catalogue of Larochelle & Larivière 2001 and the checklist of Larochelle et al. 2004) have been published until now, but keys including some native taxa have been published by Sloane (1898 and 1920; Australian taxa), Noonan (1973; world Anisodactylina genera), Moore (1977; Australian taxa), and Matthews (1980; South Australian Carabidae genera).
Prior to this revision 13 genera and 36 species of Harpalini were known from New Zealand. Following the work of Laporte de Castelnau (1867–1868), most indigenous genera and species were described before 1920 by Broun (1880–1914; 3 genera, 15 species) and Bates (1874, 1878; 3 genera, 6 species). Britton (1962, 1964a–b) and Moore (1996) provided the most recent descriptions for 2 genera, Pholeodytes and Haplanister respectively, and 6 species (including 2 in Parabaris and Syllectus). This formed the bulk of the taxonomic work on New Zealand Harpalini until now. No larval descriptions are yet available for this tribe.
If taxonomic progress has been slow until now, the collecting effort has been more intensive from the 1960s onward, so that New Zealand entomological collections and museums are now replete with Harpalini material from all areas of the country. For this reason, it seemed timely to provide a taxonomic revision for this group, one that includes descriptions and keys that take into account this new information.
The main taxonomic works that have contributed to advancing knowledge on world and New Zealand Harpalini are: Sloane (1898, key to Australian genera); Jeannel (1942, revision of France and world classification); Basilewsky (1950 and 1951, African revision); Lindroth (1968, revision of Canada, Alaska, and northern half of U.S.A.), Darlington (1968, revision of New Guinea); Habu (1973, revision of Japan); Noonan (1973, generic revision and classification of world Anisodactylina; and 1976, world catalogue of supraspecific taxa of Harpalini); Goulet (1974, revision of genus Pelmatellus); Moore (1977, key to Australian subtribes); Matthews (1980, key to South Australian genera); Moore et al. (1987, Australian catalogue); Bousquet & Larochelle (1993, Nearctic catalogue); Serrano et al. (1994, karyotypical study); Ball & Bousquet (2001, key to supraspecific taxa, North America); Larochelle & Larivière (2001, catalogue of New Zealand Carabidae); Kataev (2002a, new genus of Australian Anisodactylina); Löbl & Smetana (2003, Palaearctic catalogue).
Higher classification
The monophyly of the subfamily Harpalinae, to which belongs the tribe Harpalini, has recently been supported by molecular sequence data (Ober 2002) and larval morphology (Arndt 1998).
According to Bousquet & Larochelle (1993) the taxonomic limits of the tribe Harpalini are fairly stable although the monophyly of this taxon remains to be tested. The main contributor to the higher classification of this group was Noonan (1973, 1976) who studied the taxonomy, phylogeny, and zoogeography of the subtribe Anisodactylina and provided a synopsis of supraspecific taxa of the tribe Harpalini.
The supraspecific classification proposed by Noonan, and based on the earlier work of van Emden (1953), grouping genera into 4 subtribes (Anisodactylina, Harpalina, Pelmatellina, and Stenolophina), is generally accepted worldwide although somewhat difficult to apply in certain cases (e.g., taxonomic limits of Pelmatellina). This classification is followed here. The subtribes Harpalina, Pelmatellina, and Stenolophina still need an analysis such as provided by Noonan (1973) for Anisodactylina.
Subtribe Anisodactylina. Members of this group are distributed worldwide. About 40 genera are known (Kataev 2002a) from two genus-groups (Notiobii and Anisodactyli). According to Ball & Bousquet (2001), the Notiobii are principally in the Southern Hemisphere, showing a Gondwanan distribution pattern, whereas the Anisodactyli occur mostly in the Afrotropical and Holarctic Regions. Most New Zealand genera have the Notiobii character of the complete transverse suture between mentum and submentum. This represents the plesiomorphic state in Anisodactylina. Only the endemic Gaioxenus has the transverse suture laterally incomplete (mentum and submentum fused only medially). This is usually regarded as a character of the Anisodactyli, but exceptions have been observed by Noonan (1976) in other Southern Hemisphere taxa, e.g., within species of Anisostichus and subgenera of the Notiobia lineage, and may represent examples of parallel evolution. Noonan (1973) believed that the subtribe Anisodactylina forms a monophyletic group but he was unable to state that the group is defined on the basis of clearly apomorphic character states.
Subtribe Harpalina. Representatives of this group occur in all zoogeographical regions, mostly in tropical and temperate areas. Approximately 60 genera are known. The taxa occurring in New Zealand were introduced from Australia and the Holarctic.
Subtribe Pelmatellina. Members of this small group exhibit a Gondwanan distribution pattern in Australia, New Zealand, Andean South America, and Central America, with some taxa reaching the southwestern U.S.A. About 8 genera were described before this revision.
Pelmatellina are considered the sister group of Anisodactylina based on the shared spongily pubescent male protarsi (Noonan 1973; Goulet 1974). The current study on New Zealand taxa also agrees with Noonan (1976) on the character of the penultimate segment of the labial palpi which is bisetose (most genera) or trisetose (Kupeharpalus new genus) in Pelmatellina; not strictly bisetose as suggested by Goulet (1974). Both Noonan (1973, 1976) and Goulet (1974) indicated that pelmatelline genera differ from anisodactyline genera by the glabrous apex of the prosternal lobe. Four pelmatelline genera are now known from New Zealand, three of which share this character (Lecanomerus, Syllectus, and Hakaharpalus new genus). Kupeharpalus new genus provides the exception to this rule in having a prosternal lobe apically pubescent but in other respects sharing the characters of Lecanomerus. Further elucidation of character evolution in the Pelmatellina will have to wait until all subtribes of Harpalini are revised on a world basis.
Subtribe Stenolophina. Most species of this subtribe occur in the warm temperate and tropical regions, with 35 genera or so recorded worldwide. The morphology of New Zealand stenolophine genera, including Kiwiharpalus new genus, is consistent with the diagnostic characters provided by Noonan (1976) for this subtribe.
Noonan (1976) recorded two small endemic genera in the Australian Region (Euthenarus and Pholeodytes), to which the current revision adds the new genus Kiwiharpalus. Noonan also indicated that species of several other genera occurring in the Australian Region may be primarily centred in the Oriental Region and spreading only to the outer limit of the Australian Region or are Australian-centred taxa that may have originated from Oriental stock.
Ball & Bousquet (2001) placed the North American stenolophine genera into 2 genus-groups, Stenolophi and Polpochili. According to the literature, one important character defining the genus-group Stenolophi is the ventrally pubescent male protarsi as opposed to the absence of such pubescence in Polpochili. The study of this character in taxa indigenous to New Zealand suggests that species of Pholeodytes (endemic) and Euthenarus (not endemic) could belong to the Stenolophi. This character could not be studied in Kiwiharpalus which is known only from females. However, an Australian revision and a world reclassification and phylogeny of supra-specific taxa of Stenolophina are needed in order to uncover the true evolutionary history of this subtribe.
Geographic distribution and faunal composition
The level of endemism of the New Zealand Harpalini is 75% at the species level (42 out of 57 species) and 55% at the generic level (11 out of 20 genera). The indigenous genera Hypharpax, Lecanomerus, and Euthenarus have representatives in Australia. The genera Anisodactylus, Gnathaphanus, Notiobia, Harpalus, Egadroma, and Haplanister are adventive.
The overall distribution of New Zealand Harpalini is summarised in Table 1.
Species distributions are clearly defined and largely allopatric. Even species of a single genus, occurring in the same general areas of New Zealand are mostly allopatric within these areas (e.g., Tuiharpalus, TH–ND; Kupeharpalus, ND; Pholeodytes, NN; Hakaharpalus BR–NN–SD).
Three genera (Gaioxenus, 1 species; Parabaris, 3 species; Kupeharpalus, 3 species) are confined to the North Island. The genus Allocinopus (7 native species) occurs mostly on the North Island, except for 2 species, A. sculpticollis which is also found on the South Island, and A. latitarsis which is endemic to the Chatham Islands (CH). Two genera (Hakaharpalus, 5 species; Pholeodytes, 5 cave-dwelling species) are found only on the South Island and are restricted to the NN–SD region. Two genera are restricted to the Three Kings Islands (TH): Maoriharpalus (1 species) and Kiwiharpalus (1 species). There is no genus endemic to the Chatham Islands (CH).
Thirty-five (35) Harpalini species occur on the North Island, with 16 native species restricted to it. Thirty-one (31) species are distributed on the South Island, with 14 native species restricted to it. Only 4 indigenous species are shared between these two main islands (Allocinopus sculpticollis, Triplosarus novaezelandiae, Syllectus anomalus, and Euthenarus puncticollis). Stewart Island has no endemic taxa, but shares 2 indigenous species: Triplosarus novaezelandiae (with North Island and South Island), Euthenarus brevicollis (with South Island). Six (6) species occur on the Three Kings Islands (TH), including 4 endemics (Maoriharpalus sutherlandi, Tuiharpalus crosbyi, T. gourlayi, Kiwiharpalus townsendi), 1 adventive, and 1 indigenous species in common with the North Island (Lecanomerus sharpi). Seven (7) species occur on the Chatham Islands (CH), including 1 endemic (Allocinopus latitarsis), 2 natives in common with the South Island (Hypharpax antarcticus, Lecanomerus latimanus), and one shared with the North and South Islands (Euthenarus puncticollis), and 3 adventives. Harpalini are so far unknown from New Zealand’s subantarctic islands.
A total of fourteen (14) adventive species (about 25% of Harpalini) occur throughout New Zealand, mostly in the North Island (Map 7; especially in WN, ND, WI). The majority of adventive species probably originated from Australia apart from 2 Harpalus species and Anisodactylus binotatus (from the Palearctic Region), and Haplanister crypticus (of unknown origin).
The areas of New Zealand so far known to contain the highest diversity (Map 4) are: NN (23 species), ND (21 species), WN (17 species). The areas with the greatest number of New Zealand endemics (Map 5) are: NN (16), ND (13), BP (11).
Some Harpalini are restricted to a single area (Map 6). Currently, the areas with such species are: NN (9), ND (6), TH (4), BR (1), BP (1), CL (1), MC (1), CH (1). The South Island northwest (NN, BR) and the far north of the New Zealand (ND, TH) appear to have been the reservoirs, in geological time, of much of the genetic diversity in New Zealand Harpalini, with several species currently restricted to these regions. This trend is reflected at the generic level with Hakaharpalus occurring only in BR–NN–SD, Pholeodytes in NN, Kupeharpalus in ND, Maoriharpalus and Kiwiharpalus in TH, and Tuiharpalus in TH–ND.
Table 2 shows the genera and species shared with Australia, New Caledonia, Norfolk Island, and Lord Howe Island. Ten (10) species shared with these regions are adventive in New Zealand. Three indigenous genera (Hypharpax, Lecanomerus, Euthenarus) are shared with Australia (eastern Australian mainland and Tasmania).
Ecology, biology and dispersal power
No formal detailed study of the natural history of individual species of New Zealand Harpalini has ever been conducted although Larochelle & Larivière (2001) summarised information available from the literature, material in entomological collections, personal communications from carabid collectors, and their own personal field observations.
Native species are mostly subapterous and live within the confines of native habitats, mostly forests (especially along streams) and wet habitats, also tussock grasslands and caves (2 Syllectus, 5 Pholeodytes). The cave-dwelling species are all troglobitic, except Syllectus magnus which is troglophilous, occurring at the entrance of caves. Most Harpalini species are hygrophilous (moisture-loving) living at the surface of the soil and in leaf litter, also in caves (Syllectus, Pholeodytes), and occasionally on plants and trees. Two native species are typically found along coastal lowlands: Triplosarus novaezelandiae (on beaches and sand dunes), Allocinopus belli (coastal forests). Dispersal in native species is achieved by running over the ground; most species are moderate runners, except for the long-legged, fast-running cave species (Syllectus, Pholeodytes). In general Harpalini have relatively short legs and, sometimes, strongly reduced eyes which are indicative of strong burrowing habits.
All adventive species are macropterous and live mostly in highly modified environments (often around human dwellings), except for Haplanister crypticus which has managed to invade native forests.
The collecting period of teneral individuals suggests that Harpalini species may be either spring-breeders or summer-breeders. For most species adults are active during all months of the year, but are generally less active during cooler months.
There are no data available on the feeding preferences of Harpalini native to New Zealand. Larochelle (1990), in his review of food preferences of the Carabidae of the world, showed representatives of this tribe to be omnivorous, mostly phytophagous species. Ecomorphological adaptations providing further evidence for this feeding-type in adults and larvae have been documented by Sharova (1960, 1981), Acorn & Ball (1991), and Zetto Brandmayr et al. (1998). The mandibles of Hakaharpalus, Kiwiharpalus, Syllectus, Pholeodytes, and Maoriharpalus are unusually long among native Harpalini, which may suggest a specialised type of feeding. In addition, the strongly emarginate labrum of Maoriharpalus is reminiscent of, although not necessarily equivalent to, the condition observed in Licinini which feed on hard-bodied invertebrates, e.g., snails.
Conservation status
The Department of Conservation has responsibility for protecting and conserving New Zealand’s native plants and animals. The Department’s Species Priority Ranking System established by Molloy et al. (1994) provides criteria for scoring species according to various levels of threat, so that management and/or recovery plans can be subsequently established. A list of priority invertebrate species for conservation was established in this way by Molloy et al. (1994). McGuinness (2001) developed species profiles for species on the list, providing additional descriptive information to initiate or support key conservation actions. In addition, McGuinness (2001) added a number of invertebrates of potential conservation interest to the original list. No Harpalini species has been listed in these documents.
The Department of Conservation’s Species Ranking System is summarised in Table 3.
When the above criteria are applied, new knowledge brought forward in the present revision suggests that 24 endemic species of Harpalini (over 50% of native species) known from 10 populations or fewer may be of potential conservation concern.
All but two of these species are new to science and all species are taxonomically highly distinctive, have limited dispersal power, are often geographically localised in threatened habitats, and represented in collections by relatively few specimens collected over many decades, which may indicate rare or highly specialised species.
These species of special interest are: Allocinopus belli new species, A. bousqueti new species, A. wardi new species, Maoriharpalus sutherlandi new species, Parabaris hoarei new species, P. lesagei new species, Tuiharpalus clunieae new species, T. crosbyi new species, T. gourlayi (Britton), T. hallae new species, T. moorei new species (Anisodactylina); Hakaharpalus cavelli (Broun), H. davidsoni new species, H. maddisoni new species, H. patricki new species, H. rhodeae new species, Kupeharpalus embersoni new species, K. johnsi new species, Lecanomerus marrisi new species, Syllectus gouleti new species (Pelmatellina); Kiwiharpalus townsendi new species, Pholeodytes cerberus Britton, P. helmorei new species, P. nunni new species, and P. palmai new species (Stenolophina).