ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2006 Band/Volume: 0019 Autor(en)/Author(s): Lariviere Marie C., Larochelle Andre Artikel/Article: An overview of flat bug genera (Hemiptera, Aradidae) from New Zealand, with considerations on faunal diversification and affinities 181-214 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at An overview of flat bug genera (Hemiptera, Aradidae) from New Zealand, with considerations on faunal diversification and affinities1 M .-C . L ARIVIÈRE & A. L AROCHELLE Abstract: Nineteen genera and thirty-nine species of Aradidae have been described from New Zealand, most of which are endemic (12 genera, 38 species). An overview of all genera and an identification key to subfamilies, tribes, and genera are presented for the first time. Species included in each genus are list- ed for New Zealand. Concise generic descriptions, illustrations emphasizing key diagnostic features, colour photographs representing each genus, an overview of the most relevant literature, and notes on distribution are also given. The biology and diversification of New Zealand aradids, and their affinities with neighbouring faunas are briefly discussed. Key words: Aradidae, biogeography, Hemiptera, New Zealand, taxonomy. Introduction forests, and using their stylets to extract liq- uids from fungal hyphae associated with de- The Aradidae, also commonly referred caying wood. Many ground-dwelling species to as flat bugs or bark bugs, form a large fam- of rainforest environments are wingless – a ily of Heteroptera containing over 1,800 condition thought to have evolved several species and 210 genera worldwide (SCHUH times in the phylogeographic history of the & SLATER 1995). They are classified within group (e.g., see MONTEITH 1969b, 1982, the Pentatomomorpha and are seen by most 1997) – and have become highly modified authors as the sister group of Termitaphidi- morphologically and very strangely shaped. dae, but unlike other pentatomomorphans, It now seems amazing that before 1938 most Aradidae lack trichobothria (seta-bearing micropterous and apterous aradids were spots on the abdominal venter). thought to be only nymphs. It was MILLER Most aradid species range from 3-11 mm, (1938) who first realised the phenomenon are flattened dorsoventrally, and share the of aptery in adult Aradidae, a discovery that following diagnostic features: two-segmented opened a totally new dimension to aradid tarsi (except in some taxa); four-segmented taxonomy and accelerated the descriptive antennae; ocelli absent; elongate feeding effort in many groups. stylets (modified mandibles and maxillae) New Zealand and Australia are the only that are coiled within the clypeus when land masses so far known to harbour all withdrawn – a characteristic shared with eight recognised aradid subfamilies which, Termitaphididae – and usually broadly ex- as far as their higher classification is con- posed connexivum around the edge of rela- cerned, have remained more or less the same tively small-sized hemelytra (when present). since the comprehensive world revision of Aradids are highly cryptic animals living USINGER & MATSUDA (1959). As for tribes, either under the bark of decaying trees, or KORMILEV & FROESCHNER (1987) defined on twigs or wood debris on the floor of wet two tribes within each of the Chinamyersi- 1 The authors dedicate this paper to their friend and colleague Ernst Heiss. His encouragement toward their research, Denisia 19, zugleich Kataloge his special interest in New Zealand aradids, and his contribution to the taxonomy of the world fauna are highly in- der OÖ. Landesmuseen spirational. Neue Serie 50 (2006), 181–214 181 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at inae (Chinamyersiini and Tretocorini) and main to be described and substantial Prosympiestinae (Llaimacorini, Prosympies- amounts of unpublished information on dis- tini); all except Llaimacorini have been tribution and biology remain locked in large recorded from New Zealand. holdings of unsorted material. GROZEVA & KERZHNER (1992), in revis- Taxonomic works on New Zealand ara- ing VÁSÁRHELYI’s (1987) phylogenetic dids are scattered through the literature, the scheme, provided the most recent hypothe- depth of taxonomic treatment is unequal sis of phylogenetic relationships between between publications, and the format of tax- aradid subfamilies (Fig. 44); their classifica- onomic descriptions and the morphological tion is followed here. terms used often lack uniformity as a result of which taxa cannot be easily compared The New Zealand aradid fauna has re- within subfamilies, genera or even within cently been catalogued by LARIVIÈRE & the same publication. In addition, there is LAROCHELLE (2004), who provided a de- no identification key covering the subfami- tailed treatment of the nomenclature, geo- lies and genera known from New Zealand, graphic distribution, biology, and dispersal keys to world subfamilies do not always work power of all described genera and species — well with local taxa, and some keys to gen- the majority of which (12 out of 19 genera, era are fraught with problems (perhaps writ- 38 out of 39 species) are endemic to New ten with few specimens at hand). Zealand. Consequently, there is a need for a more The first indigenous taxon to be de- uniform and encompassing approach to the scribed from New Zealand was Ctenoneurus taxonomy of New Zealand Aradidae. This hochstetteri (MAYR 1866), followed by the de- paper aims to provide the first step towards scriptions of half a dozen or so genera and such a goal, that is: a straightforward up-to- species by WHITE (1876), MYERS & CHINA date overview of the New Zealand aradid (1928), USINGER (1943), and KORMILEV genera; an identification key to supraspecif- (1953). However, the taxonomy of New ic taxa; comparative subfamilial and generic Zealand Aradidae only really took off with descriptions; an overview of the most rele- the world revision of USINGER & MATSUDA vant literature; notes on geographic distri- (1959), who described 10 new genera and 18 bution, biology; and faunal affinities at the new species, and keyed all taxa known for generic level. the fauna at that stage. Subsequently, KOR- MILEV (1953, 1971), PENDERGRAST (1965a, 1965b, 1968), LEE & PENDERGRAST (1977), Materials and Methods KIRMAN (1985a, 1985b, 1989a, 1989b), and HEISS (1990, 1998) published additional re- Identification key visionary or descriptive papers bringing the and generic descriptions total number of taxa to that currently The starting point for the identification known. No new taxon of New Zealand Ara- key presented here was the “key to subfami- didae has been described since 1998. Korm- lies“ published by SCHUH & SLATER (1995), ilev provided much of the descriptive work which in itself was an adaptation of on a world basis following USINGER & MAT- USINGER & MATSUDA’s (1959) key. All SUDA (1959). MONTEITH (1966-1997) wrote known New Zealand species were passed the most useful contributions dealing with through SCHUH & SLATER’s key and it soon the diversification of aradid faunas in the Pa- became apparent that a number of taxa cific Southwest and Australia. would not key out to the correct subfamily. Some have claimed that the New After testing the above subfamily key Zealand aradid fauna is amongst the best and modifying it to accommodate all known known in the world. The authors are of the New Zealand genera, other keys to tribes opinion, however, after collecting Hemi- and genera were tested for all species cur- ptera in over 1,000 localities since 1992, rently known within each subfamily. In ad- and after inspecting this country’s main en- dition to USINGER & MATSUDA (1959), the tomological collections, that several taxa re- following papers containing keys were criti- 182 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at cally assessed: PENDERGRAST (1965a), which supplement and sometimes modify Aneurinae; KORMILEV (1970) and KIRMAN the world subfamily definitions derived from (1989a), Carventinae; LEE & PENDERGRAST GROZEVA & KERZHNER’s phylogeny. (1977) and MONTEITH (1997), Mezirinae; The morphological characters that var- MONTEITH (1980), Chinamyersiinae; KIR- ied most from GROZEVA & KERZHNER’s MAN (1985b), Prosympiestinae. (1992) observations were: the length of the The above process was supplemented by rostrum (Calisiinae, Mezirinae); the open- a careful analysis of published generic de- ing or closing of the rostral atrium (e.g., scriptions, together providing the basis for Carventinae, Mezirinae); the shape of the the character list utilised to write the keys pulvillus (e.g., Chinayersiinae, Mezirinae); and concise descriptions published here. the dorsal and ventral patterns of glabrous areas (in several subfamilies). The morphological characters used in this study are thought to represent the most Morphological terms diagnostic and most easily observable mor- phological features at the generic level. A An attempt has been made to use the number of non-variable character states simplest terminology available from the have been repeated between some genera to most recent aradid literature and to use facilitate their separation from world genera terms uniformly across all subfamilies. The or from potentially undescribed New main diagnostic features have been illustrat- Zealand genera. Except for the presence or ed or may be seen on the colour photo-