FNZ 63 - Auchenorrhyncha (Insecta: Hemiptera): catalogue - Introduction
Larivière, M-C; Fletcher, MJ; Larochelle, A 2010. Auchenorrhyncha (Insecta: Hemiptera): catalogue. Fauna of New Zealand 63, 232 pages.
( ISSN 0111-5383 (print), ISSN 1179-7193 (online) ; no. 63. ISBN 978-0-478-34720-3 (print), ISBN 978-0-478-34721-0 (online) ). Published 16 Jun 2010
The Auchenorrhyncha are a highly diverse group of hemimetabolous insects. They are treated here as a suborder of the Hemiptera and include the planthoppers (infraorder Fulgoromorpha), cicadas, froghoppers, spittlebugs, treehoppers, and leafhoppers (infraorder Cicadomorpha).
Auchenorrhyncha is a highly adaptable group that has evolved into a major component of the phytophagous insect fauna in most terrestrial ecosystems and adopted varied life habits on nearly all continents and islands (except Antarctica), suggesting a long evolutionary history. This insect group probably evolved since the lower Permian and over 42,000 extant species are described worldwide.
The world fauna comprises around 30 to 40 families depending on the classification being followed. Better known continental faunas such as those of North America, Europe, or Australia, include thousands of described species. Compared with these larger regions of the world, the New Zealand fauna, currently comprising 12 families, 68 genera, and 196 species, may appear relatively small, but what it lacks in size it makes up for in uniqueness, e.g., 82% of known species are endemic. From this point of view New Zealand can be regarded as a biodiversity “hot spot” for Auchenorrhyncha.
The present catalogue attempts to answer four questions most commonly asked by users of biosystematics information about a group of insects: What, where, when, and how? What Auchenorrhyncha occur in New Zealand, what is their status (e.g., endemic, native but not endemic, adventive, pest, disease vector) and what are the resources available to identify and study them? Where do species and genera occur (e.g., geographic distribution in New Zealand and overseas, habitats, dispersal abilities)? When are they active (e.g., seasonality, mating, oviposition, overwintering)? How do they live (e.g., feeding preferences, natural enemies)?
To answer these questions this catalogue brings together the available literature and collection-based information on extant taxa recorded from New Zealand’s main islands and its offshore islands. The catalogue has been written with the needs of biosystematists, identifiers, biosecurity and conservation managers, ecologists, other biologists, and members of the public in mind, hence the sections summarising for all species the geographic distribution, biology, dispersal power, and reference to available identification tools, taxonomic revisions, and natural history treatments. A species checklist, full bibliography, taxonomic index, eight appendices, species distribution maps, and primary type photographs are also provided.
Brief history of Fulgoromorpha and Cicadomorpha taxonomy in New Zealand
The first native Auchenorrhyncha described from New Zealand were the Cicadidae Amphipsalta cingulata (Fabricius, 1775) and Rhodopsalta cruentata (Fabricius, 1775). Subsequently, until about the 1930s, the majority of taxa were mostly described by European workers such as Walker (1850–1858) and White (1879), and by two New Zealand researchers Hudson (1891) and Myers (1921–1926).
Several early attempts at cataloguing the fauna were made during that same period, especially by Butler (1874), Hutton (1874, 1898, 1904), White (1879), and Kirkaldy (1909a). Such early checklists were most often straightforward compilations, but Hutton’s (1904) Index Faunae Novae Zealandiae was probably the most comprehensive and well documented, although his 1898 checklist was also very useful because it included keys to most known genera and several species. Kirkaldy’s (1909a) List of Hemiptera (excluding Sternorrhyncha) of the Maorian Subregion, with Notes on a few Species was largely based on Hutton (1904), with a few additional critical comments such as new synonymies or deletions from the fauna. Subsequently, while the New Zealand faunal inventory continued to be recorded in the various fascicles of the General Catalogue of Homoptera published by Metcalf from the 1930s to the late 1960s, there would not be another comprehensive faunal list until Wise’s (1977) A synonymic checklist of the Hexapoda of the New Zealand sub-region, the smaller orders. The latter recorded 11 families, 64 genera, and 160 species for the fauna, together with their synonyms, nomenclatural combinations, associated references, and some basic distributional information. A New Zealand checklist of Hemiptera (excluding Sternorrhyncha), listing valid names based on the 1977 list and subsequently published taxonomic changes, is continually being updated and made available on the internet (see Larivière, 2005, http://hemiptera.landcareresearch.co.nz/, New Zealand Hemiptera website). However, no complete synonymical checklist has been published since 1977.
Little taxonomic activity occurred during the 1940s and the 1950s, although Evans (1941, 1942, 1947) and Hudson (1950) added a few taxa to the fauna, in the leafhopper and cicada families respectively. The period from 1965 to 1984 was more prolific, yielding several new taxa and important taxonomic revisions, mainly due to the efforts of Fennah (1965; Delphacidae), Evans (1966; Cicadellidae sensu lato), Knight (1973–1976; Cicadellidae sensu lato), Fleming (1969, 1973, 1984; Cicadidae), Dugdale (1972; Cicadidae genera), and Dugdale & Fleming (1969, 1978; Cicadidae). Knight was, and remains, by far the most productive reviser of the New Zealand fauna, having worked on most leafhopper groups, describing new genera and approximately 50 new species.
The most recent period of active taxonomic research has occurred since 1992. Of special interest are the publications of Hamilton & Morales (1992; revision of Aphrophoridae), Larivière (1997b, 1999; revision of Cixiidae), Larivière & Hoch (1998; revision of Cixiidae), Hamilton (1999b; re-examination of Myerslopiidae genera), Emeljanov (2000; new Cixiidae genus); Larivière & Fletcher (2004; identification key to genera and species of leafhopper families); Szwedo (2004a; new Myerslopiidae species); Larivière & Fletcher (2008; Zeoliarus, new genus of Cixiidae); and Fletcher & Larivière (2009; revision of Anzygina, Cicadellidae).
The best available revisions (i.e., those containing usable keys, comparative descriptions, characters of the male genitalia, type data, comprehensive synonymies, and enough distributional information), are for the families Aphrophoridae, Cicadellidae, Cicadidae (genus Maoricicada only), and Cixiidae. However, some or part of these groups need to be re-investigated, e.g., a new key to Aphrophoridae genera is needed, and the cicadellid genera Arahura, Arawa, Horouta, Limotettix, Novothymbris, Paradorydium, Scaphetus, Matatua, and Zelopsis need additional revisionary work. Knight’s revisions of leafhopper groups and Fennah’s (1965) review of the Delphacidae were based on limited study material. A large amount of new material has accumulated in collections since the end of the 1970s and these specimens remain largely unidentified. In the Cicadidae, four out of five genera have never been revised taxonomically. It may come as a surprise to many readers that the morphological taxonomy of cicadas, such an ‘iconic’ insect group in New Zealand, is still in its infancy and that what little published information is available is patchy and scattered. Larivière et al. (2006) improved access to existing information but this cannot replace the need for a thorough taxonomic treatment of the family Cicadidae.
The Delphacidae have never been thoroughly revised, although Fennah (1965) provided preliminary insights into the New Zealand fauna. The available work on Myerslopiidae and Ulopinae (Cicadellidae) is insufficient to provide a good understanding of the fauna, mainly because very few specimens were available to earlier workers. Szwedo (2004a) made a recent effort at describing new Myerslopiidae species but ad hoc descriptions based on very small sample sizes, sometimes involving only one sex, are not always as useful as one might think, especially in groups that are in great need of overall revision.
Furthermore, so much new material has been collected and deposited in New Zealand collections over the last 30 years – one of the most dynamic insect surveying periods in New Zealand – that numerous Auchenorrhyncha taxa remain to be described even in groups worked on by previous researchers. The authors estimate that the Auchenorrhyncha fauna may reach 300 to 350 species when totally described.
Potential revisers of the fauna, whether from New Zealand or overseas, are encouraged to examine material from all the main New Zealand entomological museums and collections (see list, p. 130). They may also find it useful to note that over 65% of primary types of New Zealand Fulgoromorpha and Cicadomorpha (see type photographs, pp. 149-191) have been deposited in this country’s entomological museums and collections (see also Palma et al., 1989; Early & Gilbert, 1993; Nicholls et al., 1998), mostly in the New Zealand Arthropod Collection (NZAC). Approximately 18% of types can also be located in the Museum of Natural History (London), leaving only about 17% of types scattered among other overseas collections. The high proportion of primarily local or readily accessible type repositories means specimens can be more easily studied, making the process of revising taxa relatively less complicated for Auchenorrhyncha than for many other insect groups, which have most of their New Zealand types scattered through several overseas collections. In addition, “Virtual” collections of New Zealand types are being made available on the internet (see Larivière & Rhode, 2002, http://hemiptera.landcareresearch.co.nz/, New Zealand Hemiptera website).
Bourgoin & Campbell (2002) proposed five monophyletic groups within the Hemiptera: Sternorrhyncha (psyllids, aleyrodids, aphids, coccids), Fulgoromorpha (planthoppers), Cicadomorpha (leafhoppers, treehoppers, spittlebugs, cicadas), Heteroptera (true bugs sensu stricto), and Coleorrhyncha (moss bugs). Traditionally the Fulgoromorpha and Cicadomorpha have been referred to as Auchenorrhyncha (a suborder of Hemiptera). Several recent studies have investigated the phylogenetic relationships between the five monophyletic suborders of Hemiptera or infraorders of Auchenorrhyncha (depending on the classification followed), e.g., Campbell et al., 1995; Sorenson et al., 1995; Yoshizawa & Saigusa, 2001; Bourgoin & Campbell, 2002; Dietrich, 2002; Shcherbakov & Popov, 2002; Szwedo, 2002. Other important issues were also investigated, such as the possibility that planthoppers could be grouped under the suborder Archaeorrhyncha and other Auchenorrhyncha under Clypeorrhyncha, and whether or not Auchenorrhyncha is a monophyletic lineage.
Cryan (2005) reconstructed a molecular phylogeny of the infraorder Cicadomorpha supporting the major relationships within this group as (Membracoidea (Cicadoidea, Cercopoidea)) and compared results against previous hypotheses based on molecular and morphological data. Cryan’s work offers the most recent high-level phylogeny for Cicadomorphan families placing, as did Bourgoin & Campbell (2002), the Myerslopiidae as the sister group of the remaining Membracoidea. Cryan did not, however, test the hypothesis that Fulgoromorpha and Cicadomorpha may form a monophyletic lineage (the Auchenorrhyncha).
Urban & Cryan (2007) reviewed the main hypotheses previously put forward about the phylogeny of Fulgoroidea, both in terms of how planthopper families are related and how many families should be recognised. Their work represents the most encompassing phylogenetic hypothesis for this superfamily, so far using the greatest number of taxa and molecular markers.
Despite the ever growing body of phylogenetic work no final consensus has yet been reached on the monophyly and higher classification of Auchenorrhyncha. Pending further resolution of these matters, the traditional use of Auchenorrhyncha as a suborder or Hemiptera is retained in the present work. Table 1 follows mostly Cryan (2005) and Urban & Cryan (2007), and proposes a higher classification context in which to consider the taxa treated alphabetically in this catalogue. In Table 1, infraorders and families are arranged phylogenetically. Subfamilies and tribes of Auchenorrhyncha (excluding Cicadidae), follow Fletcher & Larivière (2001) and are listed alphabetically. The higher classification of Cicadidae follows Moulds (2005a).
Readers may find the following references useful to identify families and tribes of Auchenorrhyncha: Dietrich, 2000 (Cicadellidae), 2005 (Cicadomorpha), Fletcher & Carver, 1991 (Fulgoroidea), Fletcher et al., 1991a–b (Cercopoidea, Cicadelloidea/Membracoidea), Fletcher & Larivière, 2001 (Auchenorrhyncha), Fletcher & Stevens, 1988 (Cicadellidae), and Wilson, 2005 (Fulgoromorpha). Several links to other authoritative resources on world Auchenorrhyncha are available on the New Zealand Hemiptera website http://hemiptera.landcareresearch.co.nz/.
The New Zealand fauna is highly insular, with 41% of genera and 81% of species presently recorded as being endemic (Table 2).
The maps on pages 199–221 summarise the geographic distribution of Auchenorrhyncha species and subspecies occurring in New Zealand, based on the area codes of Crosby et al. (1976, 1998). This catalogue is the first attempt at presenting a synopsis of species distributions across all families of Fulgoromorpha and Cicadomorpha for New Zealand, based on information scattered through entomological collections as well as the literature. As a result, most species now appear to be more widely distributed than originally perceived in the literature; even well-studied species have been shown to occur in more areas of New Zealand. Nevertheless, roughly 95 native species, or 55% of the entire native fauna, are currently known from ten populations or fewer, and many of these species are known from the type locality only.
A greater number of species-group taxa (133) occur in the South Island, although 64 native taxa are actually restricted to this island. A slightly lower number of taxa (119) occur in the North Island, including 44 native taxa restricted to this island. As many as 65 taxa are shared between the two islands. For each auchenorrhynchan family the percentage of native species restricted to the North or South Island, or shared between the two islands, is shown in Table 4.
Patterns of taxonomic diversity and the number of known species-group taxa by areas of New Zealand are illustrated on Maps 4–7 (pp. 195–198). The areas so far known to contain the highest diversity (Map 4, p. 195) are, from north to south: North Island – ND (60: 47 endemics, 7 other natives, 6 adventives), AK (64: 45 endemics, 8 other natives, 11 adventives), WN (73: 63 endemics, 6 other natives, 4 adventives); South Island – NN (65: 50 endemics, 5 other natives, 10 adventives), and MC (56: 40 endemics, 5 other natives, 11 adventives). The areas so far known to harbour the highest number of New Zealand endemics (Map 5, p. 196) are: North Island – ND (47), AK and BP (45), TO (46), WN (63); South Island – NN (50), BR (45), and MC (40).
Some endemic species-group taxa are restricted to a single area of New Zealand (Map 6, p. 197). Currently, the areas known to have the greatest number of local endemics are: North Island – ND (5), WN (5); South Island – NN (4). This is closely followed by five areas of the South Island (SD, MB, MC, FD, SL), each with three known local endemics, but it is likely that the largely unexplored and unspoilt area of Fiordland (FD) is a greater reservoir of endemism than currently estimated.
The areas that include the largest number of adventive taxa (Map 7, p. 198) are: North Island – AK (11), HB (11); South Island – NN (10), MC (11). This result is hardly surprising given that these are relatively warm areas of New Zealand as well as its main trading ports or agricultural regions. Many of the adventive taxa have good dispersal abilities, some are attracted to artificial lights, and most can adapt well to living in highly or partly modified environments. Consequently a somewhat high number of adventives is also expected from neighbouring areas as can be observed on Map 7.
From the Chatham Islands seven out of 12 native species-group taxa (58% of fauna) are shared with the following New Zealand areas: North Island–South Island (1 taxon), North Island–South Island–SI (1); TH–North Island–South Island (3); KE–TH–North Island–South Island (1); TH–North Island (1). The Kermadec Islands share five out of 10 native taxa (50% of fauna) with TH–North Island–South Island (1 taxon), North Island–South Island (1), TH–North Island–South Island–CH (3). The Three Kings Islands, with 21 native taxa, have 17 of these (80% of fauna) in common with other parts of New Zealand: North Island–South Island (4 taxa); North Island–South Island–CH (1); North Island (7); North Island–CH (1); KE–North Island–South Island (1); KE–North Island–South Island–CH (1); North Island–South Island–SI (2). Auchenorrhyncha have never been recorded from New Zealand’s subantarctic islands (Antipodes, Aucklands, Bounties, Campbell Island, or Snares).
Faunal composition and affinities
Table 2 shows the number of genera and species occurring in New Zealand compared with Australia and the rest of the world. Table 3 provides a more detailed overview of the New Zealand fauna, by families and genera. The described New Zealand fauna (196 species) is about 13% the size of the known Australian fauna (around 1500 species). Currently, 15 families of Auchenorrhyncha occurring in Australia are not represented in New Zealand. In the Cicadellidae, the largest component of Cicadomorpha biodiversity in both countries, five subfamilies and 28 tribes occurring in Australia, are not represented in New Zealand.
The number of recognised adventive species in New Zealand is currently 24, or about 12% of the total Auchenorrhyncha fauna. No family is endemic to New Zealand, but all Myerslopiidae present in this country (about 70% of world species) are endemic. The three largest families in New Zealand are the Cicadellidae (78 species or 40% of the fauna), Cicadidae (34 species or 17%), and Cixiidae (26 species or 13%). These families are also well represented in Australia (see Table 2). In New Zealand, some endemic genera in the families Cicadellidae, Achilidae, Cixiidae, Delphacidae, and Derbidae are currently represented by a single species and are in need of further revision.
Most taxa shared with Australia and other parts of the world are cosmopolitan and probably introduced, except those listed in Table 5. Native taxa shared with regions neighbouring New Zealand are mostly in common with eastern continental Australia, to a lesser degree with Tasmania and Norfolk Island, and in some instances with Lord Howe Island and New Caledonia. Such faunal affinities may be indicative of a Gondwanan origin. Current data do not indicate any affinity with South America, except at the family level for Myerslopiidae. As in many parts of the world, the family Cicadellidae is taxonomically diverse and this is where most faunal affinities are observed, followed by the family Delphacidae.
At the generic level New Zealand shares 40% of its native fauna with Australia sensu lato (including Tasmania, Norfolk Island, Lord Howe Island), or 20 out of 50 native genera. At the species level this is approximately 5%. The composition of the shared native fauna does not appear to have varied much over the past several decades to 100 years.
As for the island groups in the Tasman Sea between Australia and New Zealand, Norfolk Island alone has one taxon (Kikihia: Cicadidae) with a solely New Zealand relationship. Close relationships are not generally shown with New Caledonia, except perhaps for the genus Carystoterpa (Aphrophoridae). Most taxa recorded from the Kermadec Islands are either adventive, widely distributed in the subtropical South Pacific, or (sometimes) shared with New Zealand main islands and Australia; not one species has a sole relationship with New Zealand. Only four species-group taxa (Carystoterpa subtacta, Aphrophoridae; Kikihia cutora exulis, Cicadidae; Cermada kermadecensis, Cixiidae; Ugyops (Paracona) raouli, Delphacidae) are currently recorded as being endemic to the Kermadec Islands.