FNZ 63 - Auchenorrhyncha (Insecta: Hemiptera): catalogue - Notes on families
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
Notes on families
New Zealand Auchenorrhyncha are generally diurnal and live in lowland to mountain forests and shrublands, although a number of groups are typically found in more open habitats, such as tussock grasslands, and in subalpine environments. Native species usually live within the confines of their natural habitats but some species also live in modified ecosystems and exotic tree plantations. Depending on families and genera, species can be predominantly planticolous, arboreal, or even epigean. Hostplants are known with certainty for less than 20% of taxa. The taxonomy and biology of immature stages are undescribed for the majority of taxa. Anecdotal evidence suggests that parasitic wasps, birds, predatory beetles, spiders, and mites may be among the major natural enemies of New Zealand Auchenorrhyncha. Overall, about 25% of the fauna is either brachypterous or micropterous, with active dispersal by flight being unlikely for these taxa.
Aphrophoridae. This large cosmopolitan family has relatively few representatives in New Zealand, with five genera (three natives, including one endemic) and 16 species (including 14 endemics). Also referred to as true spittlebugs, aphrophorids are xylem-feeders and usually nymphs are visible as frothy masses or “cuckoo-spit” on the stems of shrubs or small trees. However, not all species produce this froth, used by nymphs either to reduce the risk of dehydration or to deter enemies such as parasites.
Of the genera occurring on New Zealand’s main islands, Carystoterpa and Pseudaphronella are of particular interest. Carystoterpa is native genus with endemic species occurring mostly in the North Island. The habitat of many species is unknown but members of this genus are usually found on native trees and shrubs, such as Coprosma (Rubiaceae) the most likely hostplant for Carystoterpa tristis, and on surrounding exotic vegetation. Pseudaphronella jactator is the largest New Zealand aphrophorid. It is a North Island species usually found on trees and shrubs in montane to subalpine environments. Both genera are characterised by fully-winged adults and species are wide-ranging in New Zealand.
Cicadellidae. With 27 genera and 78 species distributed among ten subfamilies, the leafhoppers constitute the most diverse group of New Zealand Auchenorrhyncha. The bulk of the native fauna belongs to the subfamilies Deltocephalinae, Eupelicinae, and Tartessinae. Cicadellids occur in almost every type of habitat. New Zealand deltocephalines (e.g., Arahura, Arawa, Horouta, Limotettix) occur predominantly on low plants (very few are arboreal, e.g., Scaphetus), tartessines (Novothymbris) inhabit trees and shrubs, and eupelicines (Paradorydium) live on or very close to the ground surface. As observed elsewhere (e.g., Dolling, 1991; U.K.) brachyptery is more frequent in leafhoppers living close to the ground surface. Most leafhopper genera occurring in New Zealand feed on phloem plant sap, but the introduced Typhlocybinae feed on plant parenchyma and this is also believed to be the case for native species in this subfamily.
The economic importance of cicadellids has generated a large amount of research worldwide. In New Zealand, a handful of leafhoppers, mainly in the subfamily Typhlocybinae, have some recognised economic impact on crops. Edwardsiana froggatti (adventive) is the most important leafhopper pest in this country, feeding on pipfruit trees (especially apples) and causing a reduction in plant vigour and photosynthesis, as well as damage to fruits. Ribautiana tenerrima (adventive) feeds on raspberries, boysenberries, and blackberries, and may be a minor pest in commercial berryfruit gardens. This species may be able to transfer phytoplasmas between Rubus plants. Batracomorphus angustatus (Iassinae, possibly adventive) is a plant disease vector on tomato and potato. Anzygina dumbletoni (possibly adventive) has been found on cane fruit and strawberries, whereas Anzygina zealandica (native) is a grass species which occasionally moves into commercial orchards. Eupteryx melissae (adventive) is a potential pest of commercially grown herbs. It is known to feed on rosemary, sage, lemon balm, mint, horehound, and catmint. Nesoclutha phryne (native Deltocephalinae) and Orosius argentatus (adventive Deltocephalinae) are recognised plant disease vectors in Australia.
Native leafhopper genera of special interest. The endemic genus Arahura (Deltocephalinae: Athysanini) appears to be associated with native grass species. Additional field collecting may reveal it to contain more species and to be more widely distributed than is currently recognised.
The genus Novothymbris (Tartessinae: Thymbrini), also endemic to New Zealand, is the most diverse leafhopper genus with 16 species, and the sole representative of the subfamily Tartessinae in this country. Species distributions suggest a rich endemic fauna adapted to the montane and subalpine environments of the southern North Island mountains and the Southern Alps (South Island).
The genus Paradorydium (Eupelicinae: Paradorydiini) is also unique in that all eight New Zealand species are endemic and occur only on the South Island. One species is found on Banks Peninsula (MC), one species on Stephens Island (SD), and the remaining six species show localised allopatric distributions in the Southern Alps, mostly on each side of the Westland (WD) beech gap — a Nothofagus-impoverished area in South Westland — although P. philpotti occurs on both sides of this gap and on Stewart Island. Wing condition in Paradorydium is characterised by submacropterous coriaceous forewings and micropterous hindwings, suggesting low vagility, including an inability to fly.
The endemic monotypic genus Zelopsis is arboreal, as it is usually the case for members of the subfamily Macropsinae, with southern beech (Nothofagus) as its hostplant. The correlation is high between the distribution of Z. nothofagi and that of Nothofagus forests on New Zealand’s main islands, which may suggest related evolutionary histories and additional undescribed or extinct species in this genus.
The endemic Typhlocybinae genus Matatua (tribe Empoascini) is probably arboreal. This has been ascertained for M. maorica and is also likely for M. montivaga. The two species are allopatric on each side of the Cook Strait. These very small leafhoppers may prove more diverse and more widely distributed once their habitat is clarified through additional field collecting.
Anzygina is a native typhlocybine genus (tribe Erythroneurini) with a group of three endemic species (A. barrattae, A. ramsayi, A. toetoe) possibly representing a separate genus (see also Fletcher & Larivière, 2009) distributed on the central North Island (A. toetoe; BP, RI), on each side of the Cook Strait on the southern North Island and northern South Island (A. ramsayi; RI, BR), and in southernmost parts of the South Island (A. barrattae; SL). This apparent disjunct distribution pattern at the species level probably reflects coverage gaps in terms of field collecting and identification of material in collections and museums. The curation of these very small, easily shriveled species requires critical point drying if they are to be kept in dry storage, and also dissection of the male genitalia for accurate identification. Anzygina species have reduced forewing venation and are believed to feed on plant parenchyma. The hostplant is confirmed for A. toetoe (endemic grass toetoe or Cortaderia fulvida, Poaceae) and suspected for A. ramsayi (endemic grass tree or neinei species in the genus Dracophyllum, Epacridaceae).
The subfamily Ulopinae represents a primitive group of leafhoppers with a wide world distribution on all land masses except the Nearctic and Neotropical Regions. Ulopine leafhoppers are small, chunky insects living close to or at the surface of the ground, e.g., in the soil near the roots of plants, in leaf litter, or on low plants. They are cryptic insects, generally resembling ground debris or leaf litter fragments accumulated beneath hostplants, and have legs adapted for jumping. Vagility is low in this group with most species lacking hindwings and possessing coriaceous forewings and, consequently, incapable of flight although fully-developed hindwings have been observed in the females of some New Zealand species (Paracephaleus curtus, P. hudsoni). Szwedo (2002) hypothesised that Ulopinae might be nocturnal insects. The endemic genus Novolopa is of particular interest because it is entirely a South Island and Stewart Island group that has apparently radiated into subalpine to alpine environments. Geographical and biological knowledge on this genus is scanty. Species are thought to feed on plant phloem. Hostplants can only be hypothesised for N. falcata (Pimelea) and N. maculata (Kelleria) (both Thymelaeaceae). So far as is known, all Novolopa species lack hindwings in both sexes.
Cicadidae. Cicadas are probably among the most familiar insects due to their loud singing at the height of the New Zealand summer. Thirty-four endemic species distributed among five genera are described from this country. New Zealand cicadas are found from lowland coastal areas to the subalpine and alpine zones, in a wide range of habitats, for example coastal sand dunes, riverbeds, grasslands, scrublands, shrublands and native forests, exotic tree plantations, as well as garden and orchard hedges. Most of the forest species occur in the North Island, while the South Island is mostly characterised by a cicada fauna living in rocky open spaces. In urban environments, clapping cicadas (Amphipsalta species) can often be heard singing on garden trees, buildings, fences, and even lamp posts. The clay bank cicada (Notopsalta sericea) can also be an urban dweller in the North Island, often using any sun-warmed flat concrete surface as a singing station. The native genus Kikihia comprises 14 described species, with one species (K. convicta) known only from Norfolk Island. In New Zealand the genus has traditionally been regarded as including three sets of endemic species grouped together by Fleming (1975, 1984), mainly on the basis of habitat preferences: the “shade singers” (K. cauta, K. scutellaris), the “green foliage cicadas” (K. cutora, K. dugdalei, K. horologium, K. laneorum, K. ochrina, K. paxillulae, K. subalpina), and the “grass and scrub cicadas” (K. angusta, K. longula, K. muta, K. rosea). These groups are not supported phylogenetically (see Arensburger et al., 2004b) but they provide a convenient way to look at species that are closely similar in external morphology and quite often also in habits, ecological preferences, and/or acoustic behaviour. The black cicadas (Maoricicada species), also with 14 described species, have evolved in relation to the development of the New Zealand alpine environment where they have radiated and occupied a wide range of ecological niches.
The phylogeny and biogeography of New Zealand cicadas is a popular research topic among New Zealand and overseas scientists, especially molecular biologists. It has been the subject of several recent papers including Arensburger et al. (2004a, phylogeny and biogeography of genera; 2004b, phylogeny and dispersal patterns of Kikihia), Buckley et al. (2001a, phylogeny of Maoricicada campbelli; 2001c, origin and evolution of Maoricicada in the alpine biota; 2002, origin of New Zealand genera; 2006, evolution of Maoricicada), Buckley & Simon (2007, evolutionary radiation in Maoricicada), and Hill et al. (2009, phylogeography of Maoricicada campbelli).
These publications are readily available and results from the above studies are not repeated here, other than to indicate that 1) the closest relatives of New Zealand cicadas are from Australia and New Caledonia, 2) the evolution of the New Zealand fauna appears to have originated from multiple dispersal events (at least two) across the Tasman Sea, from Australia and possibly New Caledonia within the last 12 million years, and 3) more recently, approximately within the last five million years, speciation events led to the highly diverse genera Kikihia and Maoricicada, most likely through adaptive radiation in new habitats created by the rise of the Southern Alps.
Although the ancestors of extant taxa might have arrived via transoceanic dispersal, New Zealand cicadas have relatively short hindwings and long-distance dispersal by flight is now unlikely in most cases.
Little has been published on the biology of New Zealand cicadas apart from some basic information on singing stations, habitats, and acoustic behaviour. As in other faunas, the life of New Zealand cicadas is mostly spent in the nymphal stage, underground, and feeding on the roots of plants. Limited knowledge is available about life span, but the consensus amongst local entomologists is that at least some New Zealand species spend three to five years as nymphs, and two to four months as adults. Marris (2007) reported the following organisms as potential enemies of cicadas: parasitic wasps (cicada eggs); predatory beetles (cicada nymphs); fungal diseases (cicada nymphs and adults); kiwi birds (late instar cicada nymphs reaching the ground surface); various other birds, and spiders (cicada adults). The economic importance of cicadas is usually low, but when they occur in very large numbers significant damage may be caused by the female creating open cuts in plant tissue where eggs are laid, thus providing suitable access to boring insects and pathogens, e.g., Amphipsalta species in exotic tree plantations and fruit orchards. Pullulations (mass emergences) of cicadas can also become a major annoyance to workers in forestry and horticulture, whose activities are disrupted by loud song and repeated contact with flying insects.
The following general cicada references may be of interest to readers. Gibbs (2006) discussed the phenomenon of speciation through adaptive radiation in New Zealand cicadas. Larivière et al. (2006) provided a virtual guide to New Zealand cicadas, including identification tools and information on distribution, habitats, and songs. Marris (2007) wrote a very useful popular overview, including general information on morphological appearance, species diversity, habitats, distribution, life cycles, and enemies.
Membracidae. Only one horned treehopper species, Acanthuchus trispinifer, has so far been recorded from New Zealand and it is an Australian adventive. Membracids are usually sedentary but are capable of jumping strongly when disturbed. Adults and nymphs are often gregarious and nymphs are ant-attended. The biology and distribution of A. trispinifer in New Zealand have been studied by Eyles (1971).
Myerslopiidae. This is a small relict group of Southern Hemisphere leafhoppers, once thought to belong to the subfamily Ulopinae (Cicadellidae) but now recognised as a monophyletic lineage and a sister-group to all other Membracoidea families (Cryan, 2005). Myerslopiids are small, cryptic, ground-dwelling insects. Adults and nymphs live in leaf litter and other ground debris of high organic content forests. New Zealand taxa are characterised by large heads, spinose hindlegs, compact almost barrel-shaped bodies with cuticular extensions on head and thorax, other prominences and punctures on their coriaceous forewings, and other parts of the body that is usually encrusted with soil and litter particles offering nearly perfect camouflage with their surroundings. Functional hindwings are lacking in all species. Vestigial wing buds have been only observed in M. magna magna, the most widely distributed taxon. Consequently, active dispersal by flight is excluded for this family. The feeding strategies of Myerslopiidae are unknown (Szwedo 2004a), but they are thought to feed on fungi from the decomposing leaf and soil debris in which they live. Two endemic genera are known from New Zealand, Myerslopia (six species, two subspecies) and Pemmation (nine species, three subspecies). Of the 15 species of New Zealand Myerslopiidae, six (five Myerslopia, one Pemmation) are currently known from the type locality only.
Of the six Myerslopia species, three have a disjunct distribution in the South Island and Stewart Island. One species, M. magna, is distributed in both the North and South Islands, with the nominotypical subspecies M. magna magna occurring in the southern half of the North Island, from the Central Volcanic Plateau (or Taupo Line, approx. 39ºS) south to northwestern areas of the South Island (MB, NN, SD). The subspecies M. magna amplificata is distributed slightly further southwest in the South Island (BR), and M. magna scabrata is apparently restricted to Fiordland (FD), the south-westernmost region of the South Island. Two species are found only in the North Island and show, like their South Island congeners, a highly disjunct distribution pattern between them. Myerslopia tearohai is found in the southeast of the northern half of the North Island on Mount Te Aroha, Bay of Plenty (BP), and M. triregia is endemic to the Three Kings Islands located northwest of the North Island.
In Pemmation, three species (P. bifurca, P. terrestre, P. verrucosum) are restricted to the North Island, two species (P. asperum, P. parvum) are shared between the North and South Islands, and four species (P. insulare, P. montis, P. simile, P. townsendi) occur only in the South Island. No species is so far known from Stewart Island. Pemmation asperum is shared between both main islands, with P. asperum asperum occurring north of the Cook Strait in southernmost areas of the North Island, and P. asperum cognatum in mid-Canterbury (MC) in the South Island. Pemmation parvum shows a broad distribution from the Auckland (AK) region in the North Island, south to Westland (WD) in the South Island. Of the South Island endemics, P. townsendi is the most widely distributed with P. insulare and P. simile occupying what appears to be a subset of the distribution of P. townsendi in NN and WD respectively, although neither is known to occur in the same locality as P. townsendi.
Much more field collecting and associated taxonomic research is required before sufficient information on distribution, biology, and comparative morphology is obtained so that a reasonable understanding of the evolutionary history of this highly interesting Gondwanan group can be gained. New Zealand has 70% of the world species of Myerslopiidae, with relatives in South America (Chile) only. The genera Myerslopella Evans, 1977 (Australia) and Sagmation Hamilton, 1999 (New Caledonia, Australia) have been placed in the tribe Sagmatiini (Cicadellidae: Euacanthellinae) by Hamilton (1999b) who dismembered the subfamily Myerslopiinae based on a morphological study suggesting that superficial similarities in these genera are convergent, probably in response to similar life habits.
The disjunct distribution of Myerslopiidae in New Zealand and Chile is similar to that of Coleorrhyncha (Peloridiidae) although these also occur in Australia, Tasmania, and Lord Howe Island. According to Hamilton (1999b) there are few insect families with such Southern Hemisphere distributions, suggesting a transantarctic dispersal route available to flightless insects before the breakup of Gondwana. In the rare case of wingless genera with low number of extant species confined to Chile and New Zealand, the implication is of a very long evolutionary history of faunal elements that have persisted virtually unchanged since the Gondwanan breakup.
Achilidae. Achilid nymphs may generally be found in logs, under loose bark, or in leaf litter where they are believed to feed on fungus. Adults feed on phloem and generally lay their eggs by attaching them to woody particles in the leaf litter or to soil particles. Little is known about the biology of Agandecca annectens, the only native and endemic Achilidae known from New Zealand, but the life habits of this lowland to subalpine, forest-shrubland species may follow what is generally known for the family. Adult Agandecca annectens are macropterous, which may assist in their dispersal.
Cixiidae. Larivière (1999) revised the New Zealand fauna and provided an exhaustive account of the distribution and biology of all species. Cixiid nymphs appear to live primarily underground and feed on plant roots. Eggs are usually laid in the soil and surrounded by a waxy secretion produced by the adult female. These facts have not, however, been verified for most known species. Adults are phloem-feeders. Little is known about hostplant requirements although most New Zealand species appear to favour woody dicotyledons, a lesser number of species are associated with ferns, and very few feed on gymnosperms. Species of the genus Zeoliarus occur and feed on monocotyledons, mainly Poaceae and Phormium (Phormiaceae). Habitat preferences have been observed, for example, most New Zealand cixiids inhabit forested or bush environments, including scrublands and shrublands, from coastal lowlands to the subalpine zones. The majority of genera are found in lowland to lower mountain mixed podocarp-broadleaf habitats. The genus Aka may be closely associated with Nothofagus forests and possibly represents an older lineage. Semo is strictly a subalpine genus containing highly similar allopatric species; this may indicate relatively recent speciation. Confuga persephone is the only cave-dwelling species known from New Zealand to date. New Zealand cixiid planthoppers are mostly characterised by submacropterous to macropterous wingforms, with a tendency towards brachyptery more strongly demonstrated in the genera Aka and Chathamaka. In New Zealand the main economic importance of Cixiidae is as vectors of phytoplasma plant diseases, e.g., Zeoliarus atkinsoni, on flax (Phormium) species.
Delphacidae. Delphacids may be the most economically important planthopper family in the world because they feed on, or transmit virus diseases, to cereals, an important food source for humans globally. Adults feed on the phloem of monocots. Like the adults, nymphs roam freely. Only one species, Toya dryope, is currently recorded as being adventive to New Zealand. The native biostatus of Opiconsiva dilpa remains uncertain. However, neither of these species are recognised as plant pests or plant disease vectors in this country. Nilaparvata lugens is a vector of virus disease of rice in South-East Asia but there is no evidence of disease transmission by its New Zealand congener Nilaparvata myersi. All other species of Delphacidae are endemic to New Zealand. Wing polymorphism is displayed by some species, but most New Zealand endemics are brachypterous or, in a few cases, species may possess well-developed forewings and vestigial hindwings, e.g., Sulix. Consequently, New Zealand delphacids are thought to have low vagility by flight.
Derbidae. The nymphs of some species of Derbidae feed on fungi while most adults feed on the phloem of vascular plants. Little is known about the biology of Eocenchrea maorica, the only native and endemic derbid so far known from New Zealand. However, the life habits of this North Island, lowland-montane, forest species are expected to resemble those generally encountered in other derbids. Adults, including newly emerged individuals (tenerals), have been found on Astelia banksii (Asphodelaceae) which occurs on the forest floor of lowland forest and which may serve as a food plant. The fact that recently emerged adults have been found on this plant could provide clues about the general forest floor environment required for oviposition and nymphal development. Adults Eocenchrea maorica are macropterous, which may assist their ability to disperse.
Dictyopharidae. The family Dictyopharidae is poorly represented in the Australasian region and only one species Thanatodictya tillyardi is known from New Zealand. This species is endemic, with other Thanatodictya species occurring in Australia. Species of this genus have the head extending considerably in front of the eyes into a long process (see drawing on front cover of this contribution). Nymphs and adults feed on grass.
Flatidae. The flatid planthoppers occurring in New Zealand are Australian adventives. Their taxonomy and biology have been well studied by Fletcher (1979a, 1985, 1988, 2002). Nymphs are sessile and produce abundant wax filaments. Species are macropterous, apparently polyphagous on various plant families, feed on vascular plant phloem, and are wide-ranging in New Zealand. Anzora unicolor is of economic importance as vector of fireblight on apple and pear.
Ricaniidae. The only ricaniid planthopper occurring in New Zealand is an adventive species from Australia. The biology and economic importance in New Zealand of Scolypopa australis, the passionvine hopper, has been well studied locally, e.g., Cumber, 1966, 1967; Deitz, 1981; Hill & Steven, 1989. Biology, development, and oviposition in Australia have been described by Fletcher (1979b). This species is polyphagous and occurs on a wide range of plants in New Zealand, and is a pest of vine crops such as kiwifruit. In addition, S. australis sometimes feed on poisonous plants, e.g., tutu or Coriaria arborea, and secrete honeydew which, in times of low nectar supply, may be gathered by honey bees and incorporated into honey that is consequently poisonous to humans (Palmer-Jones et al., 1947). Scolypopa australis is a macropterous species dispersing easily and occurring in large numbers during the summer months. It is not unusual to find individual plants (native or exotic) covered by hundreds of individuals.