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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

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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- absence of well-developed paratergites on graphs provided for each genus. A few mor- the male pygophore (genital capsule), male phological features deserving further expla- genitalic characters have not been included nations are discussed here: at this early stage of the treatment of the Juga and genae. Adjacent to the clypeus New Zealand fauna because these are more dorsally are the juga (mandibular plates), taxonomically valuable at the species level. and ventrally, the genae (maxillary plates) (Fig. 1), which may or may not be produced Subfamily descriptions anteriorly to almost reach or exceed the Characters and character states were ex- apex of the clypeus. There has been some tracted from the ‘manual’ phylogeny of confusion in the literature about these struc- GROZEVA & KERZHNER (1992), which ap- tures, especially in identification keys, e.g., peared to offer the best available source the term genae should have been used in- where a set of morphological attributes had stead of mandibular plates (juga) in couplet been recorded consistently across all world 1 of SCHUH & SLATER’s (1995) key to sub- subfamilies. The present authors are aware families. that other diagnostic characters can be used Rostral atrium. When bordered by well- at the subfamily level, but they decided to developed bucculae, the rostrum may arise include only GROZEVA & KERZHNER’s (1992) from an atrium that is closed except for a characters in the basic subfamily descrip- slit-like opening (closed atrium, Fig. 8), or, tions presented here in order to test New it may arise from a wider chamber or depres- Zealand genera against GROZEVA & KERZH- sion (open atrium, Fig. 7). NER’s subfamily definitions. It soon became apparent that these subfamily definitions Rostral groove. A longitudinal furrow would need to be altered slightly to accom- running behind the rostrum along the un- modate all currently known New Zealand derside of the basal part of the head (the gu- genera. lar or ‘throat’ area).

GROZEVA & KERZHNER (1992) did not Scutellum. Usually well exposed be- indicate which taxa were seen in their study. tween the hemelytra of macropterous (Fig. These authors may have worked mostly 9), submacropterous, and brachypterous from the literature and it seems that not all species, while in micropters and apters the world genera may have been considered – scutellum becomes almost completely fused hence the New Zealand-based subfamily de- with surrounding sclerites, being only recog- scriptions included in the present paper, nisable as an elevation in the centre of the

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Figs 1-10: Generalised drawings of main diagnostic characters: (1, 2) head, dorsal view; (3) head and thorax with hemelytra reduced to wing pads, dorsal view; (4) main features of abdomen, dorsal view; (5, 6) abdomen showing placement of scent gland openings, dorsal view; (7, 8) rostral atrium on underside of head; (9) thorax and abdomen of macropterous form with well-developed scutellum, dorsal view; (10) metathoracic pleuron with well-developed evaporatory channel, lateroventral view.

184 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at mesonotum. In many apterous New Zealand included two states in their phylogenetic Carventinae, the backwardly produced me- analysis: lobelike and laminate, with the lat- dial region of the mesonotum may represent er state considered to be apomorphic. These a highly modified form of the scutellum or authors neither included the absence of pul- scutellar region. villi nor the presence of bristlelike pulvilli in their analysis, but commented in their Mesonotum and metanotum. In apter- discussion that the lack of pulvilli in Aradi- ous taxa (see photographs of many Car- nae is an apomorphic feature. Four charac- ventinae) the original segmentation may be ter states are considered in the present completely obscured by the sclerites becom- study: spatulate pulvilli not laminate apical- ing variously fused and highly modified with ly (=lobelike of above authors), spatulate tubercles, ridges, depressions (pits) or pulvilli laminate apically (=laminate of grooves. above authors), bristlelike pulvilli, and pul- Wing development. Terminology fol- villi absent. The term spatulate, also used by lows LARIVIÈRE & LAROCHELLE (2004) for MONTEITH (1997), is preferred over lobelike New Zealand Heteroptera. Macropters have because it better defines a structure that is both pairs of wings of approximately equal typically rounded or broad at the apex and length, more or less reaching apex of ab- tapered or attenuate at the base. domen. Submacropters have posterior wings Dorsal abdominal scent gland open- visibly shorter than hemelytra, and hemely- ings. These openings (Figs 4-6) are func- tra less developed than those of macropters tional and more evident in nymphs, but per- but reaching more than 2/3 of abdomen sist to various degrees in adults. Three more length. Brachypters have hemelytra reach- or less developed and equidistant paired ing 1/3-2/3 of abdomen length, the hemely- openings are located along the posterior tral membrane absent or reduced to a nar- margin of abdominal tergites III-V; in some row edging, and the posterior wings substan- groups, e.g., Mezirinae and Carventinae, the tially shorter than the hemelytra. Mi- first pair of openings are larger and strongly cropters display an extreme form of displaced posteriorly. brachyptery where hemelytra are reduced (often to simple wing pads) and posterior Tergal disc of abdomen. The abdominal wings are vestigial. segments III-VI (Fig. 4, excluding connex- ivum) are more or less fused, distinct from In aradids, the above definitions proba- previous and subsequent segments, and bly work better in terms of the relative de- forming a continuous plate called tergal velopment of each pair of wings rather than disc. of their length in relation to that of the ab- Patterns of glabrous areas domen. Apters do not have wings or, in . Smooth patches can be found on abdominal seg- many aradids, wings are not visible; small ments (dorsally (Fig. 4) and ventrally), rigid vestiges may have become fused with which have been termed glabrous areas by thoracic sclerites. USINGER & MATSUDA (1959) and shown by Parempodia and pulvilli. MONTEITH Sˇ TYS (1974) to serve as points of muscular (1997) summarised the development of the attachment to the cuticle. There can be var- nomenclature used to describe pretarsal ious arrangements of these glabrous areas. In structures. VÁSÁRHELYI (1986) applied a Mezirinae, for example, the dorsal pattern consistent terminology across all aradid sub- can be described by the formula 2:2:1, indi- families for the first time. The parempodia cating there are two glabrous areas on the consist of a pair of bristles arising from the connexivum, two midlateral glabrous areas unguitractor plate between the bases of the (next to the connexival suture), and one in- claws. The pulvilli are larger, thicker, often ner glabrous area on each side of the midline spatulate structures attached beneath each of abdominal segments III-VI. These claw (Figs 22-23). VÁSÁRHELYI (1986) glabrous areas are more visible in the nymph recognised four states for this character: than in the adult, and in the latter they are laminate, lobelike, bristlelike, and absence usually more obvious dorsally than ventral- of pulvilli. GROZEVA & KERZHNER (1992) ly.

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Figs 11-23: Generalised drawings of main diagnostic characters: (11, 12) head and thorax of some Carventinae, dorsal view; (13, 14) configuration of thoracic and first two abdominal sclerites of some Carventinae, dorsal view; (15) head with rostrum free at base, lateral view; (16) head with rostrum enclosed by bucculae at base, lateral view; (17-20) various configurations of metathoracic scent gland openings and associated evaporatory area, lateroventral view; (21) metathoracic scent gland opening with vermiculate ridges as in Tretocoris, lateroventral view; (22, 23) pretarsal pulvilli, apical view of tarsal claws and associated structures.

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Digital photographs 2(1) Rostrum arising from an open atrium Colour photographs of one representa- (Fig. 7; open area of bucculae); first dorsal abdominal scent gland opening (or scar tive of each genus were captured through a thereof) not or only slightly displaced Leica MZ-12.5 stereomicroscope, Le- posteriorly. Subfamily Aneurinae . . . . 3 icaDC500 digital camera, and the in- creased-depth-of-field syncroscopy software – Rostrum arising from a closed atrium (Fig. 8; nearly closed area of bucculae), Auto-Montage (Synoptics, U.K). Photos through a longitudinal slit-like opening – were further processed using CorelDRAW if atrium opening is widened anteriorly graphics suite version 11. As much as possi- then taxon is apterous or micropterous ble, specimens were cleared from body sur- (Acaraptera and Lissaptera (Carventinae), face incrustations (Carventinae) and cling- Woodwardiessa (Mezirinae)); first dorsal ing debris (other subfamilies) before being abdominal scent gland opening (or scar photographed (see also Note(s) under Car- thereof) displaced posteriorly to middle or ventinae and Mezirinae). hind margin of tergite IV (Fig. 6) . . . . 4 Subfamily Aneurinae Results 3(2) Scutellum poorly developed, appearing Identification key to subfamilies, as a short, posteriorly arcuate elevation in tribes, and genera the middle of the mesonotum (Fig. 32); hemelytra reduced to short lateral wing Note. Morphological characters select- pads (Fig. 3) (tergal disc of abdomen com- ed for identification are generally easy to ob- pletely exposed) (Fig. 32); connexivum serve, i.e. not requiring dissection before ex- delimited by deep longitudinal sutures amination. When a subfamily or tribe con- dorsally and ventrally; ventrites IV, V, VI tains a single genus in New Zealand, the without a distinct transverse carina along generic name is immediately given; other- anterior margin. [Habitus, Fig. 32; body wise, the identifier is referred to a subsection length, about 5 mm] ...... of the main key, a subkey to genera within a ...... Genus Aneuraptera subfamily. Additional supporting characters – Scutellum well developed, subtriangular, are given between square brackets. broadly rounded apically (Fig. 9); hemely- tra well developed, covering most of ter- 1 Genae (below juga) produced anteriorly gal disc (Fig. 9); connexivum delimited by on either side of clypeus, nearly reaching deep sutures only dorsally; ventrites IV, V, or surpassing its apex (often forming a VI with a distinct transverse carina along cleft or emarginate anterior margin of anterior margin. [Habitus, Fig. 33; body head (Fig. 1)) – genae can appear as ex- length, about 5 mm] . . . . Genus Aneurus tremely thin prolongations only clearly 4(2) Metathoracic scent gland openings visible in laterodorsal view; dorsal abdom- conspicuous, with well-developed, usually inal scent gland openings (or scars there- channel-like evaporatory area extending of) three in number, not equally devel- to lateral margin of metathoracic pleuron oped or spaced along posterior margin of (Fig. 10). [Body surface sometimes ob- tergites III, IV, V (first opening large, usu- scured by clinging dirt but not or barely ally strongly displaced posteriorly; second incrustate as in Carventinae.]. Subfamily rarely well developed; third obsolescent) Mezirinae ...... 5 (Fig. 6) ...... 2 – Metathoracic scent gland openings in- – Genae (below juga) not produced anteri- conspicuous, without well-developed orly on either side of clypeus, not nearly channel-like evaporatory area. [Body sur- reaching or surpassing its apex (never face with incrustation patterns more or forming a cleft or emarginate anterior less obscuring the integument.]. Sub- margin of head) (Fig. 2); dorsal abdomi- family Carventinae ...... 6 nal scent gland openings (or scars there- Subfamily Mezirinae of) three in number, equally developed and equally spaced along posterior margin 5(4) Rostrum arising from an open atrium of tergites III, IV, V (Fig. 5) ...... 12 (Fig. 7); hemelytra reduced to short later-

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al wing pads (Fig. 42), not extending dle into a smooth hexagonal tumescence beyond posterior margin of abdominal as above; sutures between notal sclerites tergite III (second visible); eyes stalked distinct but not very deep ...... 10 (Fig. 42). [Body hairy, especially around 10(9)Pronotum without posteriorly directed margin and on appendages; habitus, subtriangular sclerite behind collar; medi- Fig. 42; body length, about 8-10 mm] . . . al region of mesonotum produced back- ...... Genus Woodwardiessa wards as a subtriangular lobe (Fig. 34) not – Rostrum arising from a closed atrium extending as far as posterior margin of (Fig. 8); hemelytra with fully developed metanotum. [Habitus, Fig. 34; body membranes (Fig. 41), extending post- length, about 4.0-5.0 mm] ...... eriorly onto abdominal tergites VI-VII; ...... Genus Carventaptera eyes not stalked (Fig. 41). [Habitus, Fig. – Pronotum with posteriorly directed sub- 41; body length, about 6-9 mm] triangular sclerite behind collar (Fig. 14); ...... Genus Ctenoneurus medial region of mesonotum produced backwards as a subtriangular (Fig. 14) or Subfamily Carventinae subquadrate lobe extending to posterior 6(4) Rostrum arising from an open atrium margin of metanotum ...... 11 (Fig. 7; open area of bucculae); fused con- 11(10)Backward projection of medial region nexival segments II + III extending for- of mesonotum subtriangular (Figs 14, 38); wards to posterior angles of pronotum tergal disc of abdomen with segments III- (Figs 39-40) ...... 7 VI fused, without well delimited sclerites, – Rostrum arising from a closed atrium (Fig. or, partly unfused, with well delimited 8; nearly closed, slit-like area of buccu- sublateral sclerites and with inner scle- lae); fused connexival segments II + III rites partly fused with median elevation; not extending forwards to posterior angles lateral margins of metanotum flaring of pronotum ...... 8 posteriorly into more or less acutely 7(6) Meso- and metanotum fused, forming rounded projections (more produced in a smooth, uninterrupted plate (Fig. 11). males) (Fig. 14) [Habitus, Fig. 38; body [Habitus, Fig. 40; body length, about 2.5- length, about 2.5-3.5 mm] ...... 4.5 mm] ...... Genus Lissaptera ...... Genus Neocarventus – Backward projection of medial region of – Meso- and metanotum distinctly separat- mesonotum subquadrate to pentagonal ed, medial region of mesonotum produced (Fig. 37); tergal disc of abdomen with seg- backwards (Fig. 12). [Habitus, Fig. 39; ments III-VI superficially unfused, with body length, about 2.5-4.5 mm] ...... well delimited individual sublateral and ...... Genus Acaraptera inner sclerites; lateral margins of metan- 8(6) Abdominal tergites I, II completely otum not flaring posteriorly into projec- fused with metanotum and with mesono- tions. [Habitus, Fig. 37; body length, about tum at middle, produced forwards as a 2.5-3.5 mm] ...... Genus Modicarventus subtriangular lobe (Fig. 13). [Habitus, Fig. 12(1)Rostrum entirely exposed at base (not 36; body length, about 3.7-4.8 mm] . . . . bordered by well-developed bucculae) ...... Genus Leuraptera (Fig. 15); hemelytra with line of weakness – Abdominal tergites I, II separated from at level of apex of scutellum (Fig. 31), of- metanotum by a distinct suture through- ten broken off at this level. [Habitus, Fig. out, their medial regions not produced 31; body length, about 4.5-6.5 mm] . . . . forwards ...... 9 Subfamily Isoderminae, Genus Isodermus 9(8) Meso- and metanotum fused at middle – Rostrum not entirely exposed at base into a smooth hexagonal tumescence (bordered by well-developed bucculae) (Fig. 35), almost elliptical lobe in female, (Fig. 16); hemelytra, when present and extending from anterior margin of meso- well developed, without line of weakness notum almost to posterior margin of me- at level of apex of scutellum ...... 13 tanotum; sutures between notal sclerites 13(12) Metathoracic scent gland openings very deep. [Habitus, Fig. 35; body length, conspicuous with well-developed evapo- about 3.0-3.5 mm] . . . . Genus Clavaptera ratory area and channel (Figs 10, 18-20) – Meso- and metanotum not fused at mid- ...... 14

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– Metathoracic scent gland openings in- ulate (Fig. 23). [Body covered with nu- conspicuous, either small, forming a sim- merous broad, flattened granules; habitus, ple hole and without well-developed Fig. 27; body length, about 4.5-7.2 mm] . evaporatory area and channel (Fig. 17; ...... Tribe Chinamyersiini, Subfamily Aradinae), or scent gland Genus Chinamyersia opening indistinct (Subfamily Calisiinae) Subfamilies Calisiinae, Aradinae ...... 18 14(13) Metathoracic scent gland openings 18(13) Scutellum greatly enlarged, covering with a long, curved seta in middle (Figs wings and abdomen except double-edged 19-20). Subfamily Prosympiestinae .15 connexivum (Fig. 24); body tuberculate (Fig. 24), especially along margins; – Metathoracic scent gland openings with a forewings membranous, except for thick- raised ridge curved at apex (without a se- ened costal margin (Fig. 43) exposed ta) (Fig. 18) Subfamily Chinamyersiinae along basal half of scutellum. [Habitus, ...... 17 Fig. 24; body length, about 2.8-3.5 mm] . Subfamily Prosympiestinae ...... Subfamily Calisiinae, Genus Calisius 15(14) Antennae inserted on cylindrical tu- bercles (Fig. 30); metathoracic scent – Scutellum not greatly enlarged, leaving gland opening directly connecting to forewings and single-edged connexivum channel of evaporatory area (Fig. 19). broadly exposed (Fig. 25); body not tuber- [Habitus, Fig. 30; body length, about 3.0- culate; forewings of the standard hemely- 4.5 mm] ...... Genus Neadenocoris tral type (with corium and membrane). – Antennae inserted on spine-like tubercles [Habitus, Fig. 25; body length, about 5.0- (Figs 28-29); metathoracic scent gland 6.5 mm] ...... Subfamily Aradinae, opening not directly connecting to chan- Genus Aradus nel of evaporatory area (Fig. 20) . . . . 16 Overview of genera 16(15) Rostrum short, reaching or just reaching beyond posterior margin of head; Subfamily Calisiinae lateral margins of pronotum straight or Description (New Zealand): Head. nearly so, anterolateral angles subacute, Genae not large (not produced anteriorly on produced anteriorly (Fig. 29); third anten- either side of clypeus, not surpassing its nal segment distinctly shorter than other apex). Rostrum short, not reaching beyond segments. [Habitus, Fig. 29; body length, hind margin of head; bordered by bucculae about 5.5-8.0 mm] ...... at base; arising from an open atrium. Gula ...... Genus Mesadenocoris with rostral groove. Labrum free (not fused – Rostrum longer, reaching between front with clypeus). Feeding stylets coiled into coxae; lateral margins of pronotum arcu- clockwise circle. Thorax. Metathoracic ate, more or less produced laterally at mid- scent gland openings indistinct, without dle, anterolateral angles rounded, not pro- well-developed evaporatory area. Pretarsi duced anteriorly (Fig. 28); third antennal with spatulate pulvilli laminate apically. segment not distinctly shorter than other Abdomen. Posterior margin of tergites III-V segments. [Habitus, Fig. 28; body length, about 4.7-6.5 mm] . . . . Genus Adenocoris straight or nearly so. Dorsal abdominal scent gland openings (or scars thereof) equally de- Subfamily Chinamyersiinae veloped. Pattern of glabrous areas 2:1:1 17(14) Apterous. Evaporatory area of (dorsal); 2:2:1 (ventral). Paratergites of ab- metathoracic scent gland with a vermicu- dominal segment VIII of male well devel- late pattern of fine ridges (Fig. 21); tarsal oped, projecting as lobes on each side of segments 3 in number; pulvilli bristlelike pygophore. (Fig. 22). [Habitus, Fig. 26; body length, about 11 mm] ...... Tribe Tretocorini, Genus Calisius STÅL 1860 (Figs 24, 43) Genus Tretocoris Synonymy (LARIVIÈRE & LAROCHELLE 2004: 51). – Macropterous. Evaporatory area of Description (New Zealand): Body ob- metathoracic scent gland not ridged; long-oval (usually tuberculate along mar- tarsal segments 2 in number; pulvilli spat- gins), with rounded abdomen, about 2.8-3.5

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mm long. Head. Clypeus bulbous. Thorax. scars thereof) equally developed. Pattern of Scutellum greatly enlarged, apically round- glabrous areas 2:2:1 (dorsal; ventral). ed, covering wings and abdomen except Paratergites of abdominal segment VIII of connexivum (usually with three tubercles male well developed, projecting as lobes on near posterior margin of connexival seg- each side of pygophore. ments). Submacropterous; forewings almost entirely membranous, completely hidden by Genus Aradus FABRICIUS 1803 (Fig. 25) scutellum (except for thickened costal mar- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 50). gins exposed along basal half of scutellum). Description (New Zealand): Body elon- Abdomen. Connexivum broadly exposed, gate oval, about 5.0-6.5 mm long. Head. double-edged. Dorsal abdominal scent gland Clypeus bulbous. Thorax. Scutellum elon- openings (or scars thereof) three in number, gate, triangular. Submacropterous-macro- equally developed and equally spaced along pterous; hemelytra (forewings) variously de- posterior margin of tergites III, IV, V. Other veloped, completely exposed. Metathoracic characters as for subfamily. scent gland openings small, forming a sim- ple hole. Abdomen. Connexivum usually Included species: About 100 circum- broadly exposed and single-edged. Dorsal tropical species; Calisius zealandicus PENDER- abdominal scent gland openings (or scars GRAST 1968 (endemic to New Zealand). thereof) three in number, equally developed References: KORMILEV 1958 (Australian and equally spaced along posterior margin of region, key, Neotropical region, taxonomy). tergites III, IV, V. Other characters as for USINGER & MATSUDA 1959 (classification, subfamily. key to species, taxonomy, world). KORMILEV Included species: Over 200 species 1963 (Australia, key, morphology, taxono- worldwide, mostly from the Holarctic Re- my), 1966 (Australia, distribution, key, tax- gion; Aradus australis ERICHSON 1842 (na- onomy), 1967a (Australia, key to species, tive although not endemic to New Zea- South Pacific, taxonomy). PENDERGRAST land). 1968 (New Zealand, taxonomy; C. zealandi- cus). KORMILEV 1986 (keyed, taxonomy, References: USINGER & MATSUDA 1959 world). FROESCHNER 1992 (taxonomy). (classification, taxonomy, world). KORMILEV LARIVIÈRE & LAROCHELLE 2004: 51 (cata- 1966 (Australia, distribution, key, taxono- logue, New Zealand). my). PENDERGRAST 1968 (New Zealand, ta- xonomy; A. australis). LARIVIÈRE & LARO- Notes: Calisius is the only representative CHELLE 2004: 50-51 (catalogue, New Zea- of the subfamily Calisiinae in New Zealand. land). See also under Aradus. LARIVIÈRE & LAROCHELLE (2004) erroneously interpreted Notes: Because the subfamilies Aradi- the enlarged scutellum of Calisius as being nae and Calisiinae are hypothesized to be fused hemelytra. sister groups and because both subfamilies only have a single genus in New Zealand, Subfamily Aradinae the description of Aradus in this study is Description (New Zealand): Head. most directly comparative to that of Cali- Genae not large (not produced anteriorly on sius. According to USINGER & MATSUDA either side of clypeus, not surpassing its (1959), the pattern of glabrous area of Arad- apex). Rostrum long, reaching beyond hind inae is 2:1:1 (dorsal), 2:2:1 (ventral). All margin of head; bordered by bucculae at adult specimens of Aradus australis exam- base; arising from an open atrium. Gula with ined for this study had a dorsal pattern of rostral groove. Labrum free (not fused with 2:1:1, with the most inner areas barely visi- clypeus). Feeding stylets coiled into a clock- ble. The ventral pattern of 2:2:1 was more wise circle. Thorax. Metathoracic scent difficult to observe, especially the connexi- gland openings inconspicuous, without val and midlateral areas which often ap- well-developed evaporatory area. Pretarsi peared to be 1:1, perhaps because of the rel- without pulvilli. Abdomen. Posterior mar- atively small size of this species and the ven- gin of tergites III-V straight or nearly so. tral areas being less obvious in the adult Dorsal abdominal scent gland openings (or stage.

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Subfamily Chinamyersiinae Abdomen. Dorsal abdominal scent gland Description (New Zealand): Head. openings (or scars thereof) three in number, Genae not large (not produced anteriorly on equally developed and equally spaced along either side of clypeus, not surpassing its the posterior margin of tergites III, IV, V; the apex). Rostrum long, reaching beyond hind latter straight or nearly so. Abdominal spir- margin of head; bordered by bucculae at acles 5-6 ventral (2-3 lateral; 4-7, ventral; 8, base; arising from an open atrium. Gula with lateral). Pattern of glabrous areas 2:1:1 (dor- (Chinamyersia) or without (Tretocoris) ros- sal); 1:2:1 (ventral). Other characters as for tral groove. Labrum free (not fused with subfamily. clypeus). Feeding stylets coiled into a figure Included species: A single species, Treto- of eight (Chinamyersia) or into a clockwise coris grandis USINGER & MATSUDA 1959 (en- circle (Tretocoris). Thorax. Metathoracic demic to New Zealand). scent gland openings conspicuous, with References: USINGER & MATSUDA 1959 well-developed evaporatory area. Pretarsi (classification, keyed, taxonomy). MONTEI- with spatulate pulvilli not laminate apically TH 1966, 1969a (relationships), 1980 (Chinamyersia) or bristlelike pulvilli (Treto- (keyed, relationships). LARIVIÈRE & coris). Abdomen. Posterior margin of ter- LAROCHELLE 2004: 54 (catalogue, New gites III-V distinctly bent backwards (Chi- Zealand). namyersia), or, straight or nearly so (Treto- coris). Dorsal abdominal scent gland open- Tribe Chinamyersiini ings (or scars thereof) equally developed. Genus Chinamyersia USINGER 1943 Pattern of glabrous areas 1:1:1 (Chinamyer- (Fig. 27) sia) or 2:1:1 (Tretocoris) (dorsal); 1:2:1 (ven- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 54). tral). Paratergites of abdominal segment VI- Description: Body suboval to sub- II of male well developed, projecting as quadrate, attenuate anteriorly, with rounded lobes on each side of pygophore. abdomen, about 4.5-7.2 mm long, without Tribe Tretocorini short hairs but covered with numerous broad, flattened granules. Head. Eyes nor- Genus Tretocoris USINGER & MATSUDA mally developed, not stalked. Rostral groove 1959 (Fig. 26) present; feeding stylets when retracted Synonymy (LARIVIÈRE & LAROCHELLE 2004: 54). coiled into a figure of eight. Thorax. Notal Description: Body suboval to sub- lobes entire (not scalloped); pronotum with quadrate, attenuate anteriorly, with subrec- blunt anterior spine on each side of collar; tangular abdomen, about 11 mm long, cov- scutellum subtriangular to subpentagonal, ered with short hairs and surface somewhat reaching beyond posterior margin of metan- granular (not covered with numerous broad, otum. Metathoracic scent gland openings flattened granules as in Chinamyersia). present, conspicuous, with a raised ridge Head. Eyes stalked. Rostral groove absent; curved at apex (without a seta); evaporatory feeding stylets when retracted coiled into a area without a vermiculate pattern of fine simple circle. Thorax. Notal lobes scal- ridges. Macropterous. Legs without conspic- loped; pronotum without blunt spine on ei- uous spines or processes; front and middle ther side of collar; scutellum in the form of trochanters fused with femur; tarsal seg- a semicircular slightly depressed plate in the ments 2 in number; pretarsi with spatulate middle of the mesonotum, not reaching be- pulvilli not laminate apically. Abdomen. yond posterior margin of metanotum. Dorsal abdominal scent gland openings (or Metathoracic scent gland openings present, scars thereof) three in number, equally de- conspicuous, with a raised ridge curved at veloped and equally spaced along the poste- apex (without a seta); evaporatory area with rior margin of tergites III, IV, V; the latter a vermiculate pattern of fine ridges. Apter- distinctly bent backwards. Abdominal spira- ous. Legs without conspicuous spines or cles 5-6 dorsal (2-3 ventral; 4-6 dorsal; 7-8 processes; front and middle trochanters ventral). Pattern of glabrous areas 1:1:1 fused with femur; tarsal segments three in (dorsal); 1:2:1 (ventral). Other characters as number; pretarsi with bristlelike pulvilli. for subfamily.

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Included species: Two species, Chinamy- to short pads not extending beyond apex of ersia cinerea (MYERS & CHINA 1928) and C. scutellum. Metathoracic scent gland open- viridis (MYERS & CHINA 1928) (both en- ings with a long, curved seta in middle and demic to New Zealand). not directly connecting to channel of evap- oratory area, the latter strongly developed References: USINGER & MATSUDA 1959 (occupying a considerable portion of meta- (classification, keyed, taxonomy). MONTEI- pleuron and extending anteriorly and later- TH 1980 (keyed, relationships). LARIVIÈRE & ally to lateral margin). Abdomen. Tergites I, LAROCHELLE 2004: 54 (catalogue, New II well separated; III-VI fused, their posteri- Zealand). or margin indistinct. Dorsal abdominal Note: USINGER & MATSUDA (1959) re- scent glands (or scars thereof) three in num- ported “legs with unusual spines or process- ber; first opening displaced posteriorly to es“. The only unusual feature that could be middle or posterior margin of tergite IV; all observed is a granular pattern on the integu- openings equally developed. Other charac- ment of the femora, very similar to although ters as for subfamily. somewhat more spiky than that covering the remainder of the body, but nothing that Included species: Two species, Adeno- could be described as spines or processes. coris brachypterus USINGER & MATSUDA 1959 and A. spiniventris USINGER & MATSU- Subfamily Prosympiestinae DA 1959 (both endemic to New Zealand).

Description (New Zealand): Head. References: USINGER & MATSUDA 1959 Genae not large (not produced anteriorly on (classification, keyed as genus, key to either side of clypeus, not surpassing its species, taxonomy). KIRMAN 1985b (keyed). apex). Rostrum long, reaching beyond hind LARIVIÈRE & LAROCHELLE 2004: 57 (cata- margin of head; bordered by bucculae at logue, New Zealand). base; arising from an open atrium. Gula without rostral groove. Labrum fused with Genus Mesadenocoris KIRMAN 1985 clypeus. Feeding stylets coiled into an anti- (Fig. 29) clockwise circle. Thorax. Metathoracic Synonymy (LARIVIÈRE & LAROCHELLE 2004: 57). scent gland openings conspicuous, with Description: Body oval, with broadly well-developed evaporatory area. Pretarsi rounded abdomen, about 5.5-8.0 mm long. with spatulate pulvilli not laminate apically. Head. Antennae inserted on spine-like tu- Abdomen. Posterior margin of tergites III-V bercles; third segment distinctly shorter distinctly bent backwards. Dorsal abdominal than all other segments. Rostrum reaching scent gland openings (or scars thereof) or just surpassing posterior margin of head. equally developed. Pattern of glabrous areas Thorax. Pronotum with lateral margins 2:1:1 (dorsal); 2:2:1 (ventral). Paratergites straight or nearly so; anterolateral angles of abdominal segment VIII of male incon- subacute, produced anteriorly. Micropter- spicuous, not projecting as lobes on each ous; hemelytra reduced to short pads not ex- side of pygophore. tending beyond apex of scutellum. Metatho- racic scent gland openings with a long, Tribe Prosympiestini curved seta in middle and not directly con- Genus Adenocoris USINGER & MATSUDA necting to channel of evaporatory area, the 1959 (Fig. 28) latter strongly developed (occupying a con- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 57). siderable part of metapleuron and extending Description: Body suboval, about 4.7- anteriorly and laterally to lateral margin). 6.5 mm long. Head. Antennae inserted on Abdomen. Tergites I, II well separated; III- spine-like tubercles; third segment not dis- VI more or less fused, their posterior margin tinctly shorter than all other segments. Ros- mostly indistinct. Dorsal abdominal scent trum reaching between front coxae. Tho- glands (or scars thereof) three in number; rax. Pronotum with lateral margins arcuate, first opening displaced posteriorly to middle more or less produced laterally at middle; or posterior margin of tergite IV; all open- anterolateral angles rounded, not produced ings equally developed. Other characters as anteriorly. Micropterous; hemelytra reduced for subfamily.

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Fig. 24: Calisius zealandicus (Calisiinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 25: Aradus australis (Aradinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE). Fig. 26: Tretocoris grandis (Chinamyersiinae, Tretocorini), male, dorsal view. Scale bar = 1 mm. (Photo: B.E. RHODE) Fig. 27: Chinamyersia cinerea (Chinamyersiinae, Chinamyersiini), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE)

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Fig. 28: Adenocoris spiniventris (Prosympiestinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 29: Mesadenocoris robustus (Prosympiestinae), male holotype, dorsal view. Scale bar = 1 mm. (Photo: B.E. RHODE) Fig. 30: Neadenocoris spinicornis (Prosympiestinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 31: Isodermus tenuicornis (Isoderminae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE)

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Included species: A single species, References: USINGER & MATSUDA 1959 Mesadenocoris robustus KIRMAN 1985 (en- (classification, keyed as genus, key to demic to New Zealand). species, taxonomy). KIRMAN 1985b (keyed). LARIVIÈRE & LAROCHELLE 2004: 57-59 (cat- References: KIRMAN 1985b (keyed, tax- alogue, New Zealand). onomy). LARIVIÈRE & LAROCHELLE 2004: 57 (catalogue, New Zealand). Note: Examination of Prosympiestinae material from the North Island suggests ad- Note: Characters of the metathoracic ditional undescribed species that may be- scent gland openings, evaporatory area, and long to this genus or to a closely related un- abdominal tergites suggest this genus may be described genus. more closely related to Adenocoris than to Neadenocoris, although it shares a number of Subfamily Isoderminae other features with both genera. Description (New Zealand): Head. Genae not large (not produced anteriorly on Genus Neadenocoris USINGER & either side of clypeus, not surpassing its MATSUDA 1959 (Fig. 30) apex). Rostrum long, reaching beyond hind Synonymy (LARIVIÈRE & LAROCHELLE 2004: 57). margin of head; entirely exposed at base Description: Body subtriangular, with (not usually bordered by bucculae or arising broad abdomen, about 3.0-4.5 mm long. from an atrium). Gula without rostral Head. Antennae inserted on cylindrical tu- groove. Labrum fused with clypeus. Feeding bercles (sometimes slightly concave apical- stylets coiled into an anticlockwise circle. ly, but not distinctly spinous); third segment Thorax. Metathoracic scent gland openings not distinctly shorter than all other seg- conspicuous, with well-developed evapora- ments. Rostrum reaching or just surpassing tory area. Pretarsi with spatulate pulvilli posterior margin of head. Thorax. Prono- laminate apically. Abdomen. Posterior mar- tum with lateral margins straight or nearly gin of tergites III-V straight or nearly so. so; anterolateral angles subacute, produced Dorsal abdominal scent gland openings (or anteriorly. Micropterous; hemelytra reduced scars thereof) equally developed. Pattern of to short pads not extending beyond apex of glabrous areas 2:1:1 (dorsal); 2:2:1 (ventral). scutellum. Metathoracic scent gland open- Paratergites of abdominal segment VIII of ings with a long, curved seta in middle and male inconspicuous, not projecting as lobes directly connecting to channel of evapora- on each side of pygophore. tory area, the latter moderately developed Note: In Isoderminae the rostrum usual- (occupying a lesser portion of metapleuron ly arises at or near the apex of the clypeus and extending anteriorly although not later- from an anteroventral extension of its sur- ally to lateral margin). Abdomen. Tergites face. In the New Zealand species Isodermus all clearly separated; posterior margin of ter- maculosus PENDERGRAST 1965, the rostrum gites III-VI strongly sinuate on either side of arises in a more backward position, almost scent gland openings. Dorsal abdominal levelled with the antennal base, from what scent glands (or scars thereof) three in num- appears to be a small atrium bordered by ber; first opening displaced posteriorly to slightly raised bucculae. middle or posterior margin of tergite IV; all openings usually equally developed (third Genus Isodermus ERICHSON 1842 one sometimes slightly smaller). Other (Fig. 31) characters as for subfamily. Synonymy (LARIVIÈRE & LAROCHELLE 2004: 54). Included species: Six species, Neadeno- Description (New Zealand): Body elon- coris abdominalis USINGER & MATSUDA gate, almost parallel-sided, about 4.5-6.5 1959, N. acutus USINGER & MATSUDA 1959, mm long. Head. Juga well developed, reach- N. glaber USINGER & MATSUDA 1959, N. ing about middle of clypeus. Thorax. ovatus USINGER & MATSUDA 1959, N. re- Scutellum triangular. Macropterous; heme- flexus USINGER & MATSUDA 1959, and N. lytra with line of weakness at level of apex spinicornis USINGER & MATSUDA 1959 (all of scutellum, often broken off at this level. endemic to New Zealand). Prosternum forming a distinctly elevated

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plate between front coxae. Metathoracic gites III-V straight or nearly so. Dorsal ab- scent gland openings small but distinct, lo- dominal scent glands with first opening (or cated just behind middle coxae, with evapo- scar thereof) more developed than second ratory channel reaching to lateral margin of and third. Pattern of glabrous areas 2:2:1 metapleuron. Legs short; trochanters dis- (dorsal; ventral). Paratergites of abdominal tinct (not fused with femur); femora inflat- segment VIII of male well developed, pro- ed and spined. Abdomen. Dorsal abdominal jecting as lobes on each side of pygophore. scent gland openings (or scars thereof) three in number, equally developed and equally Genus Aneuraptera USINGER & spaced along posterior margin of tergites III, MATSUDA 1959 (Fig. 32) IV, V. Abdominal spiracles sublateral (al- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 48). most ventral). Other characters as for sub- Description: Body broadly oval, about 5 family. mm long. Head. Genae produced anteriorly, nearly reaching but not surpassing apex of Included species: Six species from Chile, clypeus, forming a narrowly convex anterior New Zealand, continental Australia, and margin of head (genae only visible dorsally Tasmania; Isodermus crassicornis USINGER & as extremely thin prolongations parallel to MATSUDA 1959, I. maculosus PENDERGRAST the clypeus). Juga more or less distinct on 1965 and I. tenuicornis USINGER & MATSU- either side of clypeus, only reaching near its DA 1959 (all endemic to New Zealand). base. Thorax. Scutellum in the form of a References: USINGER & MATSUDA 1959 short, posteriorly arcuate elevation in the (classification, key to species, taxonomy). middle of the mesonotum. Micropterous; PENDERGRAST 1965b (key to species, New hemelytra reduced to short lateral pads (ter- Zealand, taxonomy). KORMILEV 1966 (Aus- gal disc of abdomen completely exposed). tralia, distribution, taxonomy). HEISS 1981 Femora subcylindrical. Abdomen. Connex- (Australia, distribution, morphology, New ivum delimited by deep longitudinal sutures Zealand, South America). LARIVIÈRE & dorsally and ventrally, and with sublateral LAROCHELLE 2004: 54-55 (catalogue, New folds dorsally. Ventral surface feebly convex; Zealand). sternites IV, V, VI without a distinct trans- verse carina along anterior margin. Other Note: In Isodermus both sexes shed their characters as for subfamily. hemelytra along a line of weakness at the level of the apex of scutellum – a behaviour Included species: A single species, only found elsewhere in gerromorphan Het- Aneuraptera cimiciformis USINGER & MATSU- eroptera (SCHUH & SLATER 1995). Museum DA 1959 (endemic to New Zealand). specimens often have the apex of the heme- References: USINGER & MATSUDA 1959 lytra broken off, which may mislead some (classification, keyed, taxonomy). PENDER- people to think they represent brachypter- GRAST 1965a (taxonomy). LARIVIÈRE & ous forms. LAROCHELLE 2004: 48-49 (catalogue, New Zealand). Subfamily Aneurinae Description (New Zealand): Head. Note: In this genus, the degree of devel- Genae large (produced anteriorly on either opment of the genae on either side of the side of clypeus, nearly reaching or surpassing clypeus can be more easily observed in ven- its apex to form a cleft or emarginate anteri- trolateral view than in dorsal view. or margin of head). Rostrum short, not reach- ing beyond hind margin of head; bordered by Genus Aneurus CURTIS 1825 bucculae at base; arising from an open atri- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 49). um. Gula without rostral groove. Labrum Description: Body elongate oval, about 5 fused with clypeus. Feeding stylets coiled in- mm long. Head. Genae produced anteriorly, to an anticlockwise circle. Thorax. Metatho- reaching or surpassing apex of clypeus, form- racic scent gland openings inconspicuous, ing cleft or emarginate anterior margin of without well-developed evaporatory area. head (genae clearly visible dorsally). Juga Pretarsi with spatulate pulvilli not laminate well developed on either side of clypeus, apically. Abdomen. Posterior margin of ter- reaching about half its length. Thorax.

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Scutellum subtriangular, broadly rounded or margin of head). Rostrum short, not apically. Macropterous or submacropterous; reaching beyond hind margin of head; bor- hemelytra usually well developed (tergal disc dered by bucculae at base; arising from a of abdomen barely exposed). Femora incras- closed or open (Acaraptera, Lissaptera) atri- sate. Abdomen. Connexivum delimited by um. Gula with rostral groove. Labrum fused deep longitudinal sutures dorsally only, with- with clypeus. Feeding stylets coiled into an out sublateral folds dorsally. Ventral surface anticlockwise circle. Thorax. Metathoracic flattened, plate-like; sternites IV, V, VI with scent gland openings inconspicuous, with- a distinct transverse carina along anterior out well-developed evaporatory area. Pretar- margin. Other characters as for subfamily. si with spatulate pulvilli not laminate api- cally. Abdomen. Posterior margin of tergites Included species: Over 100 species from III-V distinctly bent backwards. Dorsal ab- all major geographic regions. dominal scent glands with first opening (or References: USINGER & MATSUDA 1959 scar thereof) more developed than second (classification, keyed, taxonomy). KORMILEV and third. Pattern of glabrous areas 2:1:1 1965 (Australia, key to species), 1966 (Aus- (dorsal, except 2:2:1 in Acaraptera and Liss- tralia, distribution, key, taxonomy), 1967b aptera); 2:2:1 (ventral, except 2:1:1 in Leu- (key to species, South America). KORMILEV raptera). Paratergites of abdominal segment & HEISS 1973 (key to species, Oriental Re- VIII of male well developed, projecting as gion). HEISS 1998 (New Zealand, revision). lobes on each side of pygophore. LARIVIÈRE & LAROCHELLE 2004: 49-50 (cat- Note: All Carventinae have the opening alogue, New Zealand). (or visible scar) of the first dorsal abdominal scent gland displaced posteriorly. As else- Subgenus Aneurodellus HEISS 1998 (Fig. 33) where in the world, New Zealand Carventi- nae have the body covered with pale incrus- Description: (HEISS 1998: 30-31). tation patterns more or less obscuring the in- Included species: Six species, Aneurus tegument. It is necessary to clear specimens (A.) brevipennis HEISS 1998, A. (A.) brouni of such incrustations before examination, WHITE 1876, A. (A.) maoricus HEISS 1998, otherwise, important diagnostic features, es- A. (A.) prominens PENDERGRAST 1965, A. pecially the shape and limits of body scle- (A.) salmoni PENDERGRAST 1965, and A. rites, may be obscured. Specimens collected (A.) zealandensis HEISS 1998 (all endemic to in ethanol can be left to dry for a few min- New Zealand). utes under a lamp and then cleaned under the microscope, before mounting, using the References: PENDERGRAST 1965a (tax- tip of a fine entomological pin or needle to onomy; as Aneurus). KORMILEV 1968a (dis- scratch off the incrustations. It is also possi- tribution, Pacific, taxonomy). HEISS 1998 ble to clean already mounted specimens us- (key to species, revision). LARIVIÈRE & ing the same method after soaking them in LAROCHELLE 2004: 49-50 (catalogue, New hot ethanol for about 30 minutes. Zealand).

Note: HEISS (1998) gave a list of charac- Group I (of USINGER & MATSUDA 1959) ters distinguishing the subgenus Aneurodel- Notes: The main diagnostic characters lus from the Palearctic Aneurus sensu stricto of this group are the closed rostral atrium and Aneurodes to which Aneurodellus ap- and the fused connexival segments II + III pears most closely related by the presence of not extending forwards to posterior angles of an area of glabrous cuticle (contergite) on pronotum. The five New Zealand genera are the inner side on the fused connexival seg- arranged alphabetically within this group ments II+III. which is one of two evolutionary lines hy- pothesized by Usinger & Matsuda (1959) for Subfamily Carventinae New Zealand genera “from the Carventus- Description: (New Zealand). Head. like ancestor … the Carventaptera, Neocar- Genae large (produced anteriorly on either ventus, Leuraptera line.“ side of clypeus, nearly reaching or surpassing its apex to form a cleft or emarginate anteri-

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Fig. 32: Aneuraptera cimiciformis (Aneurinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 33: Aneurus (Aneurodellus) zealandensis (Aneurinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 34: Carventaptera spinifera (Carventinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 35: Clavaptera ornata (Carventinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE)

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Genus Carventaptera ally almost to outer margins of eyes. Ros- USINGER & MATSUDA 1959 (Fig. 34) trum arising from a closed atrium. Thorax.

Synonymy (LARIVIÈRE & LAROCHELLE 2004: 52). Pronotum with a double-ring collar distinct- Description: Body subrectangular, about ly set off from the anterior margin, with lat- 4.0-5.0 mm long. Head. Eyes rather large in eral tubercles but lacking posteriorly direct- relation to head. Postocular spines strongly ed subtriangular sclerite. Mesonotum and developed, narrowly acutely angled, extend- metanotum distinctly separated, except at ing laterally to or slightly beyond outer mar- middle where fused into a smooth hexago- gins of eyes. Rostrum arising from a closed nal tumescence (sometimes elliptical in fe- atrium. Thorax. Pronotum with a double- male) extending from anterior margin of ring collar distinctly set off from the anterior mesonotum to posterior margin of metan- margin, with lateral tubercles but lacking otum; lateral margins of metanotum flaring posteriorly directed subtriangular sclerite. posteriorly into more or less acutely round- Mesonotum and metanotum distinctly sepa- ed projections (more developed in males). rated; medial region of mesonotum produced Sutures between notal sclerites very deep. backwards as a subtriangular lobe reaching Apterous. Abdomen. Tergites I, II separated 2/3-3/4 length (not reaching posterior mar- from metanotum and from each other by a gin) of metanotum; lateral margins of distinct suture throughout; elevated anteri- metanotum not flaring posteriorly into pro- orly; a rather thick median longitudinal ca- jections. Sutures between notal sclerites dis- rina cutting across both tergites; two antero- tinct but not deep. Apterous. Abdomen. lateral pits (one on each side) behind poste- Tergites I, II separated from metanotum by a rior margin of metanotum (as in Neocarven- distinct suture throughout; fused to each tus). Tergal disc with segments III-VI com- other and elevated anteriorly near middle. pletely fused, gradually elevated medially in- Tergal disc with segments III-VI completely to a moderate longitudinal elevation (not fused, with a distinct median longitudinal el- delimited by faint lateral margins) bearing evation with 2 median tumescent tubercles scent gland openings. Dorsal abdominal fused at middle, bearing scent gland open- scent glands (or scars thereof) three in num- ings. Dorsal abdominal scent glands (or scars ber; first opening, large, displaced posterior- thereof) three in number; first opening mod- ly to middle or posterior margin of tergite erately developed, displaced posteriorly to IV; second and third barely distinct. Fused middle or posterior margin of tergite IV; sec- connexival segments II + III not extending ond very small; third lacking. Fused connex- forwards to posterior angles of pronotum. ival segments II + III not extending forwards Only spiracles 3-7 lateral and visible from to posterior angles of pronotum. Only spira- above. Pattern of glabrous areas 2:1:1 (dor- cles 5-7 lateral and visible from above. Pat- sal); 2:2:1 (ventral). Other characters as for terns of glabrous areas 2:1:1 (dorsal); 2:2: subfamily. 1(ventral). Other characters as for subfamily. Included species: A single species, Included species: A single species, Car- Clavaptera ornata KIRMAN 1985 (endemic to ventaptera spinifera USINGER & MATSUDA New Zealand). 1959 (endemic to New Zealand). References: KIRMAN 1985a (taxonomy), References: USINGER & MATSUDA 1959 1989a (keyed). LARIVIÈRE & LAROCHELLE (classification, keyed, taxonomy). KIRMAN 2004: 52 (catalogue, New Zealand). 1989a (keyed). LARIVIÈRE & LAROCHELLE 2004: 52 (catalogue, New Zealand). Genus Leuraptera USINGER & MATSUDA 1959 (Fig. 36)

Genus Clavaptera KIRMAN 1985 Synonymy (LARIVIÈRE & LAROCHELLE 2004: 52). (Fig. 35) Description: Body oval, about 3.7-4.8 Synonymy (LARIVIÈRE & LAROCHELLE 2004: 52). mm long. Head. Eyes very small in relation Description: Body subrectangular, about to head. Postocular spines strongly devel- 3.0-3.5 mm long. Head. Eyes rather small in oped, acutely angled, extending laterally to relation to head. Postocular spines slightly or slightly beyond outer margins of eyes. developed, obtusely angled, extending later- Rostrum arising from a closed atrium. Tho-

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Fig. 36: Leuraptera zealandica (Carventinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 37: Modicarventus wisei (Carventinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 38: Neocarventus angulatus (Carventinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 39: Acaraptera wisei (Carventinae) showing some body surface incrustations, female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE)

200 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at rax. Pronotum with a double-ring collar dis- mesonotum produced backwards as a sub- tinctly set off from the anterior margin, with quadrate to pentagonal lobe reaching poste- lateral tubercles but lacking posteriorly di- rior margin of metanotum; lateral margins of rected subtriangular sclerite. Mesonotum metanotum not flaring posteriorly into pro- and metanotum distinctly separated; medial jections as in Clavaptera or Neocarventus. region of mesonotum and metanotum sepa- Sutures between notal sclerites distinct but rated in two by forward-extended subtrian- not deep. Apterous. Abdomen. Tergites I, II gular median lobe of abdominal base; lateral separated from metanotum by distinct su- margins of metanotum not flaring posterior- ture throughout; mostly fused to each other, ly into projections. Sutures between notal but with trace of a faint suture between the sclerites distinct but not deep. Apterous. two; a tubercle-bearing median longitudinal Abdomen. Tergites I, II separated from groove cutting across both sclerites; two an- metanotum and from each other by a dis- terolateral slits (as opposed to pits in tinct suture laterally only. Tergal disc with Clavaptera and Neocarventus) (one on each segments III-VI completely fused, usually side) behind posterior margin of metan- without (sometimes with faint) median lon- otum. Tergal disc with segments III-VI su- gitudinal elevation (not delimited by faint perficially unfused, with well delimited indi- lateral margins) bearing scent gland open- vidual sublateral and inner sclerites (4 sub- ings. Dorsal abdominal scent glands (or lateral and 4 inner sclerites); only slightly scars thereof) three in number; first opening elevated at middle into short longitudinal moderately developed, displaced posteriorly elevation bearing scent gland openings. to middle or posterior margin of tergite IV; Dorsal abdominal scent gland openings (or second very small; third evanescent. Fused scars thereof) three in number; first opening connexival segments II + III not extending large, displaced posteriorly slightly behind forwards to posterior angles of pronotum. middle of segment IV; second very small; Only spiracles 5-7 lateral and visible from third lacking. Fused connexival segments II above. Pattern of glabrous areas noticeable + III not extending forwards to posterior an- although sometimes faint, 2:1:1 (dorsal; gles of pronotum. Only spiracles 1, 5-7 lat- ventral). Other characters as for subfamily. eral and visible from above. Pattern of glabrous areas 2:1:1 (dorsal); 2:2:1 (ventral). Included species: Two species, Leu- Other characters as for subfamily. raptera yakasi HEISS 1990 and L. zealandica USINGER & MATSUDA 1959 (both endemic Included species: A single species, Modi- to New Zealand). carventus wisei KIRMAN 1989 (endemic to New Zealand). References: USINGER & MATSUDA 1959 (classification, keyed, taxonomy). KIRMAN References: KIRMAN 1989a (keyed, tax- 1989a (keyed). HEISS 1990 (taxonomy; L. onomy). LARIVIÈRE & LAROCHELLE 2004: 53 yakasi). LARIVIÈRE & LAROCHELLE 2004: 52 (catalogue, New Zealand). (catalogue, New Zealand). Genus Neocarventus USINGER & Genus Modicarventus KIRMAN 1989 MATSUDA 1959 (Fig. 38)

(Fig. 37) Synonymy (LARIVIÈRE & LAROCHELLE 2004: 53). Synonymy (LARIVIÈRE & LAROCHELLE 2004: 53). Description: Body subovate (N. angula- Description: Body broadly pear-shaped tus) or subrectangular (N. uncus), about 2.5- to subrectangular, about 2.5-3.5 mm long. 3.5 mm long. Head. Eyes rather large in re- Head. Eyes rather small in relation to head. lation to head. Postocular spines lacking or Postocular spines barely developed, forming barely developed, forming narrow postocular narrowly rounded postocular swellings. Ros- swellings (N. angulatus) or slightly devel- trum arising from a closed atrium. Thorax. oped, obtusely angled, nearly extending to Pronotum with a double-ring collar distinct- outer margins of eyes (N. uncus). Rostrum ly set off from the anterior margin, with lat- arising from a closed atrium. Thorax. Prono- eral tubercles and posteriorly directed sub- tum with a double-ring collar distinctly set triangular sclerite. Mesonotum and metan- off from the anterior margin, with lateral tu- otum distinctly separated; medial region of bercles and posteriorly directed subtriangular

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sclerite. Mesonotum and metanotum dis- pronotum. The two New Zealand genera tinctly separated; medial region of mesono- (Acaraptera and Lissaptera) are arranged al- tum produced backwards into triangular (N. phabetically within this group which is the angulatus) or subtriangular (N. uncus) lobe second evolutionary line hypothesized by reaching posterior margin of metanotum; USINGER & MATSUDA (1959) to include lateral margins of metanotum flaring posteri- New Zealand genera “the Acaraptera-Che- orly into more or less acutely rounded pro- lonaptera line [lapsus]“ [= Acaraptera-Biroana jections (more developed in males and in N. line (Monteith pers. comm.)] and thought uncus). Sutures between notal sclerites dis- to be further removed from a Carventus-like tinct but not deep. Apterous. Abdomen. ancestor than Group I. Further comments Tergites I, II separated from metanotum by a on Group II may be found in MONTEITH distinct suture throughout; separated from (1967). The genus Nesiaptera (islands of each other laterally only (N. angulatus) or Samoa and Fiji) is the closest relative of throughout (N. uncus); elevated anteriorly; Acaraptera and Lissaptera. a rather thick median longitudinal carina cutting across both tergites (as in Clava- Genus Acaraptera USINGER & MATSUDA ptera); two anterolateral pits (as in Clava- 1959 (Fig. 39) ptera) (one on each side) behind posterior Synonymy (LARIVIÈRE & LAROCHELLE 2004: 51). margin of metanotum. Tergal disc with seg- Description: Body pear-shaped, about ments III-VI completely fused (N. angulatus) 2.5-4.5 mm long. Head. Eyes rather large in or in part superficially unfused (N. uncus) relation to head. Postocular spines moder- with well delimited sublateral sclerites and ately to barely developed, usually quadrate, with inner sclerites partly fused with median extending laterally almost to outer margins elevation, the latter gradually elevated to- of eyes, or, sometimes forming narrowly wards middle (delimited by lateral margins), rounded postocular swellings. Rostrum aris- bearing scent gland openings. Dorsal abdom- ing from an open atrium. Thorax. Pronotum inal scent gland openings (or scars thereof) with a single-ring collar barely set off from three in number; first opening moderately the anterior margin, without tubercles or developed, displaced posteriorly to middle or posteriorly directed subtriangular sclerite. posterior margin of tergite IV; second very Mesonotum and metanotum distinctly sepa- small; third lacking. Fused connexival seg- rated; medial region of mesonotum pro- ments II + III not extending forwards to pos- duced backwards as a subtriangular lobe terior angles of pronotum. Only spiracles 5-7 reaching into anterior 1/3 of metanotum; lateral and visible from above. Pattern of lateral margins of metanotum not flaring glabrous areas 2:1:1 (dorsal); 2:2:1 (ventral). posteriorly into projections. Sutures be- Other characters as for subfamily. tween notal sclerites distinct but not deep. Apterous. Abdomen. Tergites I, II separated Included species: Two species, Neocar- from metanotum by a distinct suture ventus angulatus USINGER & MATSUDA 1959 throughout; fused to each other (A. myersi) and N. uncus KIRMAN 1989 (both endemic or fused only laterally, separated by a dis- to New Zealand). tinct suture medially (A. waipouensis). Ter-

References: USINGER & MATSUDA 1959 gal disc with segments III-VI completely (classification, keyed, taxonomy). KIRMAN fused, with a distinct median longitudinal 1989a (keyed), 1989b (redescription, taxon- elevation (delimited by faint lateral mar- omy). LARIVIÈRE & LAROCHELLE 2004: 53 gins) bearing scent gland openings. Dorsal (catalogue, New Zealand). abdominal scent gland openings (or scars thereof) three in number; first opening dis- Note: Neocarventus is morphologically tinct, large, displaced posteriorly to middle closer to Clavaptera than to Modicarventus. or posterior margin of tergite IV; second smaller; third evanescent or lacking. Fused Group II (of USINGER & MATSUDA 1959) connexival segments II + III extending for- Notes: The main diagnostic characters wards to posterior angles of pronotum. Spir- of this group are the open rostral atrium and acles lateral, all visible from above. Pattern the fused connexival segments II + III ex- of glabrous areas 2:2:1 (dorsal; ventral). tending forwards to posterior angles of Other characters as for subfamily.

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Included species: Five described species margin of tergite IV; second very small; – Lord Howe Island (2), New Zealand (2), third evanescent or lacking. Fused connexi- Solomon Islands (1); also two undescribed val segments II + III extending forwards to species from Norfolk Island (1) and New posterior angles of pronotum. Spiracles lat- Caledonia (1) (Monteith pers. comm.); eral, all visible from above. Pattern of Acaraptera myersi USINGER & MATSUDA glabrous areas 2:2:1 (dorsal; ventral). Other 1959 and A. waipouensis HEISS 1990 (both characters as for subfamily. endemic to New Zealand). Included species: Two species from Lord References: USINGER & MATSUDA 1959 Howe Island and New Zealand; Lissaptera (classification, keyed, taxonomy). KOR- completa (USINGER & MATSUDA 1959) (en- MILEV 1966 (Australia, distribution, New demic to New Zealand). Zealand, sexual dimorphism, taxonomy). References: USINGER & MATSUDA 1959 MONTEITH 1967 (distribution, morphology). (classification, keyed, taxonomy; as sub- KORMILEV 1968b (distribution, taxonomy). genus of Acaraptera). KORMILEV 1966 (Aus- HEISS 1990 (taxonomy). LARIVIÈRE & tralia, distribution, New Zealand, sexual di- LAROCHELLE 2004: 51 (catalogue, New morphism, taxonomy). KORMILEV 1968b Zealand). (distribution, taxonomy). LARIVIÈRE & Note: USINGER & MATSUDA (1959) de- LAROCHELLE 2004: 52-53 (catalogue, New scribed three subgenera, Acaraptera, Liss- Zealand). aptera, and Nesiaptera, that were elevated to Subfamily Mezirinae generic status by KORMILEV & FROESCHNER (1987). Description (New Zealand): Head. Genae large (produced anteriorly on either Genus Lissaptera USINGER & MATSUDA side of clypeus, nearly reaching or surpassing 1959 (Fig. 40) its apex to form a cleft or emarginate anteri-

Synonymy (LARIVIÈRE & LAROCHELLE 2004: 52). or margin on head). Rostrum short, not Description: Body pear-shaped, about reaching beyond hind margin of head 2.5-4.5 mm long. Head. Eyes rather large in (Woodwardiessa), or, sometimes longer, relation to head. Postocular spines barely to reaching beyond hind margin (Ctenoneu- moderately developed, usually forming nar- rus); bordered by bucculae at base; arising rowly rounded postocular swellings, or, from a closed (Ctenoneurus) or open (Wood- sometimes more quadrate, extending later- wardiessa) atrium. Gula with rostral groove. ally almost to outer margins of eyes. Ros- Labrum fused with clypeus. Feeding stylets trum arising from an open atrium. Thorax. coiled into an anticlockwise circle. Thorax. Pronotum with a single-ring collar barely set Metathoracic scent gland openings conspic- off from the anterior margin, without tuber- uous, with well-developed evaporatory area. cles or posteriorly directed subtriangular Pretarsi with spatulate pulvilli not laminate sclerite. Mesonotum and metanotum com- apically. Abdomen. Posterior margin of ter- pletely fused, forming a smooth uninterrupt- gites III-V distinctly bent backwards. Dorsal ed plate; lateral margins of metanotum not abdominal scent glands with first opening flaring posteriorly into projections. Sutures (or scar thereof) more developed than sec- between notal sclerites distinct but not ond and third. Pattern of glabrous areas deep. Apterous. Abdomen. Tergites I, II 2:2:1 (dorsal, ventral). Paratergites of ab- partly separated from metanotum by a very dominal segment VIII of male well devel- short suture medially; completely fused to oped, projecting as lobes on each side of each other. Tergal disc with segments III-VI pygophore. completely fused, with or without a slight Notes: Mezirinae lack the body surface median longitudinal elevation (not delimit- incrustations found in Carventinae, but ed by faint lateral margins) bearing scent they are often covered with clinging debris. gland openings. Dorsal abdominal scent This is especially true of ground-dwelling gland openings (or scars thereof) three in apterous or micropterous species (e.g., number; first opening distinct, large, dis- Woodwardiessa quadrata), which need to be placed posteriorly to middle or posterior cleaned before examination. Specimens col-

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Fig. 40: Lissaptera completa (Carventinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 41: Ctenoneurus hochstetteri (Mezirinae), male, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 42: Woodwardiessa quadrata (Mezirinae), female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE) Fig. 43: Calisius zealandicus (Calisiinae) with extended wings, female, dorsal view. Scale bar = 1 mm. (Photo: M.-C. LARIVIÈRE)

204 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at lected in ethanol can be brushed under the References: USINGER & MATSUDA 1959 microscope, before mounting, using a fine (classification, keyed, taxonomy). KOR- art brush that has been cut very short at the MILEV 1971 (key to genera, key to species, tip (see MONTEITH 1997). It is also possible Oriental region, South Pacific, revision). to clean already mounted specimens with LEE & PENDERGRAST 1977 (key to species, the same method after soaking them in hot revision, New Zealand). MONTEITH 1997 ethanol for about 30 minutes. (Australia, keyed, revision). LARIVIÈRE & LAROCHELLE 2004: 55-56 (catalogue, New Genus Ctenoneurus BERGROTH 1887 Zealand). (Fig. 41) SINGER Synonymy (LARIVIÈRE & LAROCHELLE 2004: 55). Genus Woodwardiessa U & Description (New Zealand): Body elon- MATSUDA 1959 (Fig. 42) gate, subparallel, nearly glabrous, about 6.0- Synonymy (LARIVIÈRE & LAROCHELLE 2004: 56). 9.0 mm long. Head. Eyes normally devel- Description: Body subquadrate, attenu- oped, not stalked as in Woodwardiessa. Ros- ate anteriorly, pubescent (especially dorsal- trum arising fom a closed atrium (nearly ly, on appendages, and along body margin), closed slit-like area of bucculae), extending about 8.0-10.0 mm long. Head. Eyes stalked. Rostrum arising from an open atri- beyond limits of rostral groove and some- um (open area of bucculae) not extending times onto prosternum. Thorax. Anterolat- beyond limits of rostral groove or onto eral angles of pronotum obtusely rounded, prosternum. Thorax. Anterolateral angles not produced anteriorly. Scutellum subtri- of pronotum acutely rounded, produced an- angular, as long or a little longer than teriorly. Scutellum in the form of a semicir- pronotum, with prominent spines at basal cular plate in the middle of the mesonotum, angles extending forwards over posterior about ? as long as pronotum, without for- margin of pronotum. Macropterous; hemely- wardly extended spines as in Ctenoneurus, tra with fully developed membranes, ex- but with distinct lobes at basal angles. Mi- tending posteriorly onto abdominal tergites cropterous; hemelytra in the form of short VI-VII. Metathoracic scent gland openings pads on either side on mesonotum, not ex- conspicuous, located just behind middle tending beyond posterior margin of abdomi- coxae, with evaporatory channel reaching nal tergite III (second visible). Metatho- to lateral margin of metapleuron. Legs with racic scent gland openings conspicuous, lo- trochanters distinct (not fused to femur); cated just behind middle coxae, with evapo- femora moderately inflated, without spines. ratory channel reaching to lateral margin of Abdomen: Tergal disc mostly concealed be- metapleuron. Legs with trochanters distinct neath wings, flattish, smooth and glabrous. (not fused to femur); femora moderately in- Dorsal abdominal scent gland openings (or flated, without spines. Abdomen. Tergal scars thereof) three in number, not equally disc exposed, rather convex, rugose and pit- developed or spaced along the posterior ted, with erect hairs. Dorsal abdominal margin of tergites III, IV, V (first opening scent gland openings (or scars thereof) three large, usually strongly displaced posteriorly in number, not equally developed or spaced to middle or posterior margin of tergite IV; along the posterior margin of tergites III, IV, second rarely well developed; third obsoles- V (first opening large, usually strongly dis- cent). Sternites IV, V, VI with a more or less placed posteriorly to middle or posterior developed transverse carina along anterior margin of tergite IV; second rarely well de- margin. Other characters as for subfamily. veloped; third obsolescent). Sternites IV, V, VI without a transverse carina along anteri- Included species: Over 40 species from or margin. Other characters as for subfamily. the Neotropical, Ethiopian, Oriental, and Australian biogeographic regions; Ctenoneu- Included species: A single species, Woodwardiessa quadrata USINGER & MATSU- rus hochstetteri (MAYR 1866), C. myersi KOR- DA 1959 (endemic to New Zealand). MILEV 1953, C. pendergrasti KORMILEV 1971, and C. setosus LEE & PENDERGRAST 1977 References: USINGER & MATSUDA 1959 (all endemic to New Zealand). (classification, keyed, taxonomy). KOR-

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Fig. 44: Phylogeny of Aradidae subfamilies from GROZEVA & KERZHNER (1992).

MILEV 1971 (keyed). MONTEITH 1997 (Aus- tralia, keyed, revision). LARIVIÈRE & LAROCHELLE 2004: 56-57 (catalogue, New Zealand).

Note: KORMILEV (1964) described Woodwardiessa australiensis (= Neophloeobia australiensis) from Australia but transferred it as the type species of Scirrhocoris KOR- MILEV 1965, which was later synonymised with Neophloeobia USINGER & MATSUDA 1959, by MONTEITH (1997).

Discussion

Considerations on faunal diversification and affinities Table 1 summarises the distribution and some of the most remarkable biological at- tributes of the 19 genera of New Zealand Aradidae in taxonomic sequence by subfam- ilies, in a manner similar to MONTEITH (1997) for Australian Mezirinae. The geo- graphic regions of New Zealand used to record distributions are shown in Figure 45 and represent broadly accepted large-scale faunistic areas. Of the two main islands of New Zealand, the North Island has the most diverse fauna, with 11 out of 19 genera recorded as being endemic to it; only the Fig. 45: The New Zealand subregion with areas of the North and South Islands: ND, genus Neadenocoris (Prosympiestinae) is Northland (northernmost North Island); N, northern North Island; S, southern North currently known from the South Island on- Island; NW, northwestern South Island; NE, northeastern South Island; SW, southwestern South Island; SE, southeastern South Island; OT, Otago. ly. Before discussing the diversification and

206 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at affinities of New Zealand’s aradid fauna any cene; some smaller more disjunct pockets al- further, it is necessary to provide some back- so occur northward but usually mixed with ground information into the country’s natu- or in close proximity to broadleaf-podocarp ral environment. forests. Northland, the northernmost region of the North Island, is acknowledged as an The New Zealand environment. New old and geologically quite stable faunistic Zealand lies in the south-west of the South centre (WARDLE 1991). In addition, temper- Pacific about 1,500 km west of its nearest ate rainforests have remained extensive neighbour Australia. The country is long (c. there for long stretches of time, even during 1600 km) and narrow, with its two main is- the last glaciation (as recently as 10,000 lands spanning many latitudes (c. 34°-46°S) years ago) (WARDLE 1991) when the remain- and characterised by a complex physiogra- der of New Zealand was mostly covered by phy including several mountain ranges shrub- and grasslands with pockets of forests reaching great heights in the Southern Alps in the lowlands (podocarp-broadleaf in the (South Island) and dominated by volcanic North, beech in the South) and low-growing land forms on the North Island central alpine vegetation at higher elevations. plateau. Consequently, several climatic zones are recognised (KIRKPATRICK 1999): Figure 45 also shows the South Island to the northern half of the North Island has be divided into eastern and western regions warm humid summers and mild winters delimited roughly by the main divide of the (slightly cooler in the east) with an annual Southern Alps, and into northern and rainfall of 1000-2000 mm; in the central southern regions at about 43°-44°S because North Island (especially the central vol- this latitudinal range corresponds to a tran- canic plateau) conditions vary with altitude sitional faunistic zone observed in other in- and exposure, but many areas have a high sect groups (e.g., other Hemiptera and rainfall mountain climate; the southern half ground-beetles (Coleoptera: Carabidae)). of the North Island has warm summers and The Otago region (including Central Otago mild winters with high rainfall (1300-2000 and Otago Lakes) is a well-documented cen- mm) in the west, and very warm summers tre of biodiversity forested by Nothofagus un- and moderate winters with lower rainfall til, according to MCGLONE (1988), fire per- (600-1000 mm) along the east coast; much manently destroyed much of the forest of the eastern South Island has warm sum- about 2,500 years ago. Nowadays much of mers with cool winters (with frequent frosts this area is covered by native grasslands and occasional snow) and annual rainfall of (mainly tussock grasslands), open shrub- 500-1500 mm (slightly less inland in the ex- and scrublands, as well as relatively large treme south); the Otago region has very hot tracts of adventive pastures. summers, cold winters, and semiarid condi- Overall, New Zealand’s vegetation cov- tions; much of the South Island west coast er has been altered considerably from its has a climate that varies little throughout pre-human condition, from 75 % forest cov- the year, with small temperature ranges and er to only about 23 %, but WARDLE (1991) high rainfall increasing rapidly eastward estimated that “native vegetation, albeit of- with altitude (mountain climate conditions ten greatly modified, still covers about 60 % prevail in the Southern Alps). of New Zealand; more than a quarter of it In Figure 45, the North Island is divided protected in national parks and other re- into northern and southern regions along serves.“ the “Taupo-line“ – a biogeographic feature Faunal diversification and affinities. referring to a band across the central North Speculating too much on the diversification Island at approximately 39°-39°30’S (GIBBS of the New Zealand fauna without more de- 1989), which roughly runs across a line of tailed taxonomic and phylogenetic analyses old volcanoes, with Mount Ruapehu as its could be hazardous but it is possible, based focal point on the central volcanic plateau. on extant distribution patterns and current This “line“ defines more or less the current knowledge of the biology of New Zealand northern limit of the re-establishment of taxa, to identify some trends and formulate beech (Nothofagus) forests since the Holo- hypotheses for further investigation.

207

© Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at OTHER REGIONS OTHER

Samoa, Ethiopian, Oriental

SOUTH AMERICA SOUTH NEW CALEDONIA NEW

Presence of a genus in the series geo- nifer“), Sub= Subcorticolous (under tree Lord Howe I Howe Lord Norfolk I Norfolk

apterous, Mi=Micropterous, Ma=macropterous, Tasmania

st North Island (Northland and/or Three Kings Islands). Mainland

Chatham Is. Chatham

Stewart Island Stewart

S Island (SW) Island S

S Island (Otago) Island S

S Island (SE) Island S

S Island (NE) Island S S Island (NW) Island S

NEW ZEALAND AUSTRALIA

N Island (southern) Island N

N Island (northern) Island N

both pairs of wings are membranous, somewhat reduced and covered by the enlarged scutellum. Is Kings Three

Calisius Microhabitat

Rainforest type Rainforest

Altitudinal range Altitudinal

Flight ability Flight Endemism En Ma Lo-Mo BrPo, Be Sub En Ap Lo-Mo BrPo Supra En Ap Lo-Mo BrPo Supra En Ap Lo BrPo Supra En Mi Lo BrPo Supra En Ap Lo BrPo Supra En Ap Lo BrPo Supra En Mi Lo BrPo Supra En Ap Lo-Mo BrPo, Be Supra En Mi Lo-Sa Be Supra En Mi Lo-Mo BrPo, Be Supra? Na Ma Lo-Sa BrPo, Be Sub X X Micronesian, Melanesian, W Na sMa* Lo-Sa BrPo, Be Supra? X X X X X Circumtropical Na Ap Lo BrPo Supra X Na sMa-Ma Lo-Mo BrPo, Be Sub X X X X Cosmopolitan Na Ma Lo-Sa Be Sub X X Chile Na Ap Lo-Mo BrPo Supra X X X Solomons ) En sMa-Ma Lo-Sa BrPo, Be Sub En Mi Lo BrPo Sub Aradus Calisius Tretocoris Lissaptera Isodermus Clavaptera Leuraptera Acaraptera Adenocoris Ctenoneurus Aneuraptera Chinamyersia Neadenocoris Neocarventus Aneurodellus Carventaptera Modicarventus ( Mesadenocoris : Summary of geographic distribution and biology the 19 genera New Zealand Aradidae in taxonomic sequence by Subfamilies. Aneurus Tretocorini Prosympiestini I Group II Group Mezirinae Woodwardiessa Calisiinae Aradinae Chinamyersiinae Chinamyersiini Prosympiestinae Isoderminae Aneurinae Carventinae graphic regions is indicated by a solid cell (New Zealand) or X (elsewhere); black cells indicate taxa found only in northernmo sMa= Submacropterous, Lo=lowland, Mo=montane, Sa=Subalpine, Be=beech (Nothofagus), Br=broadleaf („evergreen“), Po=podocarp („co bark), Supra=Supracorticolous (on exterior of bark wood debris). *In Tab. 1 Tab. Entries for other columns are as follows: En=endemic (restricted) to New Zealand, Na=native, but not endemic Zealand; A=

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From what is known of their geographic served at the family level. Suprageneric distribution, habits, and affinities with taxa groupings show a paucity of macropterous of neighbouring land masses (Table 1, see al- taxa compared with continental faunas or so LARIVIÈRE & LAROCHELLE 2004), there is less isolated island faunas: the Prosympiesti- little doubt New Zealand aradids have had a nae are known from micropterous taxa only; long evolutionary history closely linked to the known Carventinae are all apterous; the that of rainforest habitats, in particular, old relative ancestry of Tretocoris (apterous Tre- Gondwanan elements such as podocarps tocorini) and Chinamyersia (macropterous (‘conifers’) and Nothofagus (beech) that oc- Chinamyersiini) cannot be established – the curred widely and were shared with other same situation applies to the relationship parts of Gondwana. between the micropterous Aneuraptera and submacropterous to macropterous Aneurus Aptery. Wing loss in Aradidae as an (Aneurodellus) in Aneurinae; Woodwardiessa evolutionary novelty related to life in rain- (micropterous Mezirinae) may be a relict forest microhabitats has been discussed by Gondwanan taxon. Consequently, wing loss MONTEITH (1969b, 1982). Submacropterous may have been well on its way in the an- and macropterous taxa would have evolved cient podocarp-dominated forests of Gond- as specialist inhabitants of the thin space wana before New Zealand separated from it (subcortical space) beneath the loose bark (80 mya). of dead trees in relatively open and drier forests where flight ability is required to Carventinae and Mezirinae. Northland, colonise suitable habitats. On the other or the northernmost region of the North Is- hand, mostly ground-dwelling, micropterous land, belonged to a complex Tertiary island or apterous inhabitants of rainforests would arc system stretching from the direction of have evolved a more sedentary life on the the islands of New Guinea and New Cale- outside of bark. In such habitats, logs and donia. The Carventinae and Mezirinae of other wood debris on the moist forest floor, Northland are of particular interest because do not shed their bark easily, usually remain five monotypic micropterous or apterous to decay for long periods of time (without genera are restricted to this region (Mesade- the subcortical space), and are in close prox- nocoris – Prosympiestinae; Aneuraptera – Aneurinae; Clavaptera, Modicarventus – Car- imity to each other. MONTEITH (1982) fur- ther elaborated that “freed from the con- ventinae; Woodwardiessa – Mezirinae). straints of flight ability and flat shape, and Acaraptera is also an interesting apterous confronted with the need for camouflage, carventine genus that was previously made we thus see the evolution in rainforests up of three subgenera (Acaraptera, Lissaptera around the world of Aradidae that have lost and Nesiaptera) until these were given full their wings and assumed the grotesque, tu- generic status by KORMILEV & FROESCHNER (1987). Acaraptera sensu stricto occurs on bercular form of bark-mimics. The end the North Island of New Zealand, Lord point of this faunal evolution appears to be Howe Island, Norfolk Island (Monteith a stable situation where regional aradid fau- pers. comm.), the Solomon Islands, and nas consist of a complement of normal, New Caledonia, while Nesiaptera is found macropterous, subcortical species coexisting on the islands of Samoa and Fiji. Lissaptera with a complement of totally apterous, bark- is shared between the Three Kings Islands exterior [supracorticolous] species.“ (60 km northwest of the North Island) and This interpretation is supported by ob- Lord Howe Island. Acaraptera sensu lato has servations in New Zealand (Table 1) where been hypothesized by MONTEITH (1982) to extant taxa from putatively derived subfam- represent an ancient lineage dating from the ilies are mostly ground-dwelling, supracorti- more extensive lands of the Melanesian Arc colous, apterous or sometimes micropterous region (Eocene to late Miocene), possibly species occurring in regions that have sup- related to the genus Biroana, which occurs ported rainforest habitats over long periods at high altitudes in the New Guinea moun- of geological time. In the extant New tains. MONTEITH (1982) also regarded Zealand fauna, however, the full comple- Acaraptera sensu lato to be largely a moun- ment of wing conditions can only be ob- tain element on the islands where it occurs;

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this interpretation is not supported by New of the closely related genera Acaraptera, Lis- Zealand data (see Table 1). saptera, and Nesiaptera may provide further evidence of this Southwest Pacific link. Woodwardiessa is a micropterous mezirine that may be a relict taxon with ex- Chinamyersiinae. Members of this sub- tant, more derived relatives in Australia and family are restricted to Australia, New New Caledonia. Its morphological affinities Zealand, New Caledonia, and Vanuatu. and hypothesized phylogenetic relationship MONTEITH (1980) hypothesized that extant to a rainforest group of seven apterous Aus- Chinamyersiinae constitute an ancient tralian genera closely related to the New ‘relict’ group with two distinct lineages: the Caledonian genus Phloeobia, led MONTEITH macropterous Chinamyersiini containing (1997) to suggest an Australian plate-New one species of Gnostocoris (Vanuatu and Zealand-New Caledonia link maintained New Caledonia) and two species of Chi- since before the separation of these land namyersia (New Zealand; subcorticolous); masses, and that the Australian and New the apterous Tretocorini including three Caledonian genera probably evolved from species of Kumaressa (eastern Australia) and Mezira-like ancestors through forms equiva- one species of Tretocoris (New Zealand; lent to Woodwardiessa. supracorticolous). The two tribes have de- The world Carventinae and Mezirinae veloped fundamental morphological differ- are closely related and have evolved in trop- ences, mainly in the stylet coiling and pre- ical and subtropical environments into over tarsal pulvilli, but also differentiated body 60 genera and 250 species and 120 genera forms correlated with winged or wingless and 1,100 species, respectively. In these sub- conditions and life habits. Insufficient infor- families aptery is thought to be a derived mation is available to establish with certain- condition that has arisen many times. The ty the relative ancestry of these lineages. absence of macropterous Carventinae from MONTEITH (1969a) thought Kumaressa and New Zealand and the extent of morpholog- Tretocoris must have shared a common ical evolution of apterous genera, mainly ev- macropterous ancestor and that the sympa- idenced by thoracic and abdominal sclerite try of Tretocoris and Chinamyersia in New modification and fusion, together with their Zealand might offer some clues to that ef- obvious low dispersal ability, provide sup- fect, but he also suggested that the morphol- port for the hypothesis that much of their ogy of the Chinamyersiini might have fol- evolutionary history may have occurred in lowed a completely different evolutionary situ, possibly in very restricted geographical path from that of the Tretocorini. If Treto- areas before the breakup of New Zealand coris and Chinamyersia were sympatric dur- from Gondwana (80 mya) or thereafter. Fur- ing New Zealand’s geological history, this ther support for this idea may be offered by pattern might not be so easily observed to- the low representation of Mezirinae in New day. The authors are not suggesting that this Zealand and the fact that the only endemic could not be the case, but further field col- genus is represented by the ‘relict’ mi- lecting is needed to establish whether Chi- cropterous species Woodwardiessa quadrata. namyersia cinerea and Tretocoris grandis do in fact co-occur in the same North Island lo- MONTEITH (1982) already suggested calities and habitats. that the mostly apterous carventine faunas of Australia, New Guinea, New Zealand and Prosympiestinae. KORMILEV & New Caledonia represent an ancient ele- FROESCHNER (1987) recognised two tribes in ment of the Southwest Pacific. In addition, this subfamily, the Llaimocorini for Llaimo- the Northland region of New Zealand, coris penai (type species of a macropterous where most apterous Carventinae are found, monotypic genus from Chile), and the retained pockets of subtropical rainforests Prosympiestini (New Zealand, Australia during much of New Zealand’s geological (including Tasmania), and New Guinea). history, which developed more recently into Currently, Adenocoris (one species), Mesade- warm-temperate rainforests that were in ex- nocoris (one species), and Neadenocoris (six istence even during the last glacial phase. species) are known from New Zealand, and As discussed above, the extant distributions Prosympiestus (four species) from eastern

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Australia. Consequently, this subfamily can found in New Zealand; although rarely col- be regarded as having a typical eastern lected it has been found in a number of Gondwana distribution even though none places across the North Island and in the of the extant genera and species are shared northern half of the South Island. Very lit- between South America, Australia and New tle is known about the biology of Calisiinae Zealand. According to CASSIS & GROSS in general. In the Palearctic Region Calisius (2002) the Australian Prosympiestus species has been observed feeding on the phloem of are all macropterous and most likely subcor- living trees with natural cracks in the bark ticolous. The New Zealand Prosympiesti- (see personal communication by Kerzhner nae, on the other hand, are all micropterous in LARIVIÈRE & LAROCHELLE 2004: 51). In and supracorticolous. Adenocoris and Australia, species are thought to be able to Mesadenocoris occur on the North Island on- fly (CASSIS & GROSS 2002). Monteith (pers. ly. Neadenocoris, the most diversified and comm.) reports that they live mostly in possibly more recently evolved genus, is cur- small cracks and crevices on the outside of rently known from the South Island only rough bark of dead trees and dead logs, and where species are distributed allopatrically their dappled colour pattern gives them from lowland to subalpine beech forest habi- good camouflage in this situation – hun- tats. The cold-adapted beech forests of New dreds of specimens representing numerous Zealand did occur on Gondwana before its species have been collected in this situation breakup but they are a more recent Gond- in Australia and New Caledonia by spraying wanan element than the podocarp forests. the trunks of dead trees with pyrethrin. The current distribution of the three New In New Zealand, the habitat of C. Zealand genera and of species of Neadena- zealandicus is unclear. Collecting records coris also suggests that diversification in the currently show the species to have been latter might have been facilitated by chang- equally found on trees and shrubs as on fall- ing climatic conditions and alternate expan- en logs or on the ground. Records of the sion and constriction of forest habitats dur- species coming to artificial lights (LARIVIÈRE ing the Pleistocene. & LAROCHELLE 2004) may provide indirect Isoderminae. This subfamily includes a evidence of flight. single genus, Isodermus, with a classic east- Aradinae and Aneurinae. Distribution- ern Gondwanan distribution in Chile (one al and biological data of interest on Aradi- species), New Zealand (three species) and nae and Aneurinae are noted in Table 1. southeast Australia, including Tasmania Aradus australis is the only species of Aradi- (two species); no species is shared between nae occurring in New Zealand; it is also dis- these areas. Isodermus species are subcorti- tributed in Australia and New Caledonia. colous and macropterous in New Zealand, The New Zealand Aneurinae are perhaps mostly distributed south of the ‘Taupo-line’ more interesting. Aneuraptera and Aneu- on the North Island and throughout most rodellus (subgenus of Aneurus) are both en- areas of the South Island. Their range corre- demic to New Zealand; the former includes sponds well with the extant distribution of a single supracorticolous micropterous beech forests. species from the old pordocarp-broadleaf Calisiinae. This subfamily is known forests of Northland; the latter contains six from six genera and about 100 species subcorticolous submacropterous to macro- worldwide. The genus Calisius countains pterous species in the podocarp-broadleaf most species and occurs circumtropically. In and beech forests of the North and South Is- the neighbourhood of New Zealand it is lands. found mostly in New Guinea, Australia (southern and eastern continental, Tasma- nia, Norfolk Island), Melanesia (including New Caledonia) and Polynesia. Thirteen endemic species are found in Australia, with the majority in the forests of Queensland. A single endemic species, C. zealandicus, is

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Acknowledgements References

The authors wish to thank Robert CASSIS G. & G.F. GROSS (2002): Hemiptera: Het- Hoare, Rosa Henderson and Anne Austin eroptera (Pentatomomorpha). — In: HOUSTON W.W.K. & G.V. MAYNARD (Eds), Zoological Cat- (Landcare Research, New Zealand) for re- alogue of Australia. Vol. 27 3B. CSIRO Aus- viewing the manuscript. The authors are al- tralia, Melbourne: xiv + 1-737.

so grateful to Birgit Rhode (Landcare Re- FROESCHNER R.C. (1992): The flat bug genus Cali- search, New Zealand) for her help in ob- siopsis CHAMPION: A review with descriptions taining important scientific papers and for of three new species, including one from Do- the preparation of two colour photographs. minican Republic amber (Heteroptera: Aradi- dae). — Proc. Biol. Soc. Wash. 105 (1): 32-39. Thanks are also extended to Geoff Montei- th (Brisbane Museum, Queensland, Aus- GIBBS G.W. (1989): Local or global? Biogeography of some primitive Lepidoptera in New tralia) for personal communications, for his Zealand. — New Zealand Journal of Zoology permission to use a table format similar to 16: 689-698. that used in Monteith (1997: 162, Fig. 69) GOEL S.C. & C.W. SCHAEFFER (1970): The structure of to summarise biology and distribution data, the pulvillus and its taxonomic value in the and for writing several highly insteresting land Heteroptera (Hemiptera). — Ann. Ento- papers on Australian and South Pacific Ara- mol. Soc. Amer. 63: 307-313. didae – without which there would be very GROZEVA S.M. & I.M. KERZHNER (1992): On the phy- limited context to discuss the New Zealand logeny of aradid subfamilies (Heteroptera: Aradidae). — Acta Zool. Hung. 38 (3-4): 199- fauna. Finally, many thanks to Ernst Heiss 205. (Tiroler Landesmuseum, Innsbruck, Aus- HEISS E. (1981): On Isodermus planus ERICHSON, tria) for his collegial support and encourage- 1842, and a new species from Tasmania (Het- ment toward the authors’ heteropterological eroptera, Aradidae). — Entomofauna 2 (22): studies and his special interest in New 247-262.

Zealand aradids. This project was supported HEISS E. (1990): New apterous Carventinae from in part by funding from the Foundation for New Zealand (Heteroptera: Aradidae). — J. Research, Science and Technology (con- NY Entomol. Soc. 98 (4): 393-401. tract number C09X0501), New Zealand. HEISS E. (1998): Review of the genus Aneurus from New Zealand with description of three new species (Heteroptera, Aradidae). — New Zusammenfassung Zealand Journal of Zoology 25: 29-42. Neunzehn Gattungen und 39 Arten der KIRKPATRICK R. (1999): Bateman contemporary atlas New Zealand. — David Bateman Limited, Aradidae wurden aus Neuseeland beschrie- Auckland, New Zealand: 1-160. ben, die meisten sind endemisch (12 Gat- KIRMAN M. (1985a): Clavaptera ornata n. gen. et tungen, 38 Arten). Die Arbeit gibt einen sp., a new genus and species of Carventinae Überblick aller Gattungen und enthält erst- (Hemiptera: Heteroptera: Aradidae) from mals einen Bestimmungsschlüssel der Northland, New Zealand. — New Zealand Unterfamilien, Triben und Gattungen. Die Journal of Zoology 12: 125-129. neuseeländischen Arten werden aufgelistet. KIRMAN M. (1985b): A new genus and species of Kurze Gattungsdiagnosen, Zeichnungen der Prosympiestinae (Hexapoda: Hemiptera: Ara- didae) from Northland, New Zealand. — wichtigsten Merkmale, Farbphotos aller Records of the Auckland Institute and Muse- Gattungen, ein Überblick zur wichtigsten um 22: 77-83. Literatur und Angaben zur Verbreitung wer- KIRMAN M. (1989a): A new genus and species of den präsentiert. Die Biologie und Mannig- Carventinae (Hexapoda: Hemiptera: Aradi- faltigkeit der Aradiden Neuseelands und ih- dae) from Northland, New Zealand. — re Beziehungen zu benachbarten Faunen Records of the Auckland Institute & Museum werden diskutiert. 26: 25-32. KIRMAN M. (1989b): A redescription of the genus Neocarventus (Hexapoda: Hemiptera: Aradi- dae) and a description of a new species from Northland, New Zealand. — Records of the Auckland Institute & Museum 26: 33-38.

KORMILEV N.A. (1953): Notes on Aradidae from the Eastern Hemisphere. — Verh. Naturf. Ges. Basel 64: 333-346.

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KORMILEV N.A. (1958): Notes on Aradidae in the MCGLONE M.S. (1988): New Zealand. — In: HUNTLEY U.S. National Museum. 1. Subfamily Calisi- B. & T. WEBB III (Eds), Handbook of Vegetation inae. — Proc. US Nat. Mus. 109 (3413): 209- Science 7. Vegetation History. Kluwer Acade- 222. mics Publishers, Dordrecht: 557-599.

KORMILEV N.A. (1963): On some Calisiinae in the MILLER N.C.E. (1938): A new subfamily of British Museum (Nat. Hist.) (Hemiptera-Het- Malaysian Dysodiidae (Rhynchota). — Ann. eroptera: Aradidae). — Ann. Mag. Nat. Hist. Mag. Nat. Hist. 11(1): 498-510. (13) 5, (1962): 601-607. MONTEITH G.B. (1966): A new genus of Chinamyer-

KORMILEV N.A. (1964): New genera and species of siinae (Heteroptera: Aradidae) from Aus- Queensland Aradidae (Hemiptera: Hetero- tralia, with notes on its relationships and ptera). — J. Entomol. Soc. Queensland 3: 42- male genitalia. — J. Entomol. Soc. Queens- 47. land 5: 46-50.

KORMILEV N.A. (1965): Notes on Australian Aradi- MONTEITH G.B. (1967): A revision and redescription dae (Hemiptera: Heteroptera) with descrip- of the genus Glyptoaptera KORMILEV tions of new genera and species. — Proc. R. (Hemiptera: Aradidae). — Proc. R. Entomol. Soc. Queensl. 77 (3): 11-35. Soc. London (B) 36: 50-60.

MONTEITH G.B. (1969a): The relationship of Kuma- KORMILEV N.A. (1966): Aradidae in the South Aus- tralian Museum, Adelaide (Hemiptera-Het- ressa MONTEITH and Tretocoris USINGER & MAT- eroptera). — Rec. South Austr. Mus. 15: 275- SUDA, with a new species of Kumaressa (Hemiptera: Aradidae: Chinamyersiinae). — 307. Proc. R. Soc. Queensl. 81: 75-82. KORMILEV N.A. (1967a): Notes on Australian Aradi- MONTEITH G.B. (1969b): A remarkable case of alary dae (Hemiptera: Heteroptera) with descrip- dimorphism in the Aradidae (Hemiptera) tions of new species of Calisius STÅL and Glo- with a generic synonymy and a new species. chocoris USINGER & MATSUDA. — Proc. R. Soc. — J. Aust. Entomol. Soc. 8: 87-94. Queensl. 79: 71-78. MONTEITH G.B. (1980): Relationships of the genera KORMILEV N.A. (1967b): On some Aradidae from of Chinamyersiinae, with description of a Brasil, Argentina and Laos (Hemiptera: Het- relict species from the mountains of North eroptera). — Opusc. Zool. 100: 1-10. Queensland (Hemiptera: Heteroptera: Aradi- KORMILEV N.A. (1968a): Aradidae in the Bishop Mu- dae). — Pac. Ins. 21 (1979): 275-285. seum, Honolulu, II (Hemiptera: Heteroptera). MONTEITH G.B. (1982): Biogeography of the New — Pac. Ins. 10 (2): 249-260. Guinea Aradidae (Heteroptera). — Mono- KORMILEV N.A. (1968b): Aradidae in the Bishop Mu- graphiae Biologicae, The Hague 42: 645-657. seum, Honolulu, III (Hemiptera: Heteroptera). MONTEITH G.B. (1997): Revision of the Australian — Pac. Ins. 10 (3-4): 575-597. flat bugs of the subfamily Mezirinae (Insecta: KORMILEV N.A. (1970): Aradidae in the Bishop Mu- Hemiptera: Aradidae). — Mem. Queensland seum, Honolulu V. (Supplement) (Hemiptera- Mus. 41 (1): 1-169. Heteroptera). — Pac. Ins. 12 (4): 701-722. MYERS J.G. & W.E. CHINA (1928): A list of New KORMILEV N.A. (1971): Mezirinae of the Oriental Zealand Heteroptera with the description of Region and South Pacific (Hemiptera-Het- a remarkable green aradid representing a eroptera: Aradidae). — Pac. Ins., Monograph new genus. — Ann. Mag. Nat. Hist. (10) 1 (3): 26: 1-165. 377-394.

KORMILEV N.A. (1986): Two new genera and seven PENDERGRAST J.G. (1965a): The Aradidae of New new species of Aradidae (Heteroptera). — J. Zealand (Hemiptera, Heteroptera) I. Introduc- NY Entomol. Soc. 94: 249-261. tion; the Aneurinae. — Trans. Roy. Soc. New Zealand, Zoology 6 (5): 53-63. KORMILEV N.A. & R.C. FROESCHNER (1987): Flat Bugs of the World: A synonymic list (Heteroptera: PENDERGRAST J.G. (1965b): The Aradidae of New Aradidae). — Entomography 5: 1-246. Zealand (Hemiptera, Heteroptera) II. The Iso- derminae. — Trans. Roy. Soc. New Zealand, KORMILEV N.A. & E. HEISS (1973): A new species of Zoology 6 (23): 235-243. the genus Aneurus CURTIS 1825, from Nepal PENDERGRAST J.G. (1968): The Aradidae of New (Hemiptera: Aradidae). — Entomol. Rec. 85: Zealand (Hemiptera, Heteroptera) III. The 141-146, plate 10. Aradinae and Calisiinae. — Trans. Roy. Soc. LARIVIÈRE M.-C. & A. LAROCHELLE (2004): Heteroptera New Zealand, Zoology 10 (10): 81-88. (Insecta: Hemiptera): catalogue. — Fauna of SCHUH R.T. (1976): Pretarsal structure in the Miri- New Zealand 50: 1-330. dae (Hemiptera) with a cladistic analysis of re- LEE C.E. & J.G. PENDERGRAST (1977): The Aradidae of lationships within the family. — Am. Mus. New Zealand (Hemiptera: Heteroptera) IV. Novitates 2601: 1-39. The Mezirinae, with a consideration of the SCHUH R.T. & J.A. SLATER (1995): True Bugs of the genitalia. — New Zealand Journal of Zoology world (Hemiptera: Heteroptera). Classifica- 4 (2): 167-175. tion and natural history. — Cornell University MAYR G.L. (1866): Diagnosen neuer Hemipteren III. Press, Ithaca & London: 1-336 [Aradidae, pp. — Verh. Zool.-Bot. Ges. Wien 16: 361-366. 208-214].

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Sˇ TYS P. (1974): Morphological and taxonomic notes on the Aneurinae, with descriptions of Aneu- rus (Iralunelus subgen. n.) gallicus sp. n. from France, and a world list of species (Het- eroptera: Aradidae). — Acta Entomol. Bohe- moslov. 71: 86-104.

USINGER R.L. (1943): A new name for a New Zealand aradid. — Pan-Pacific Entomol. 19: 74.

USINGER R.L. & R. MATSUDA (1959): Classification of the Aradidae. — British Museum, London: 1- 410.

VÁSÁRHELYI T. (1986): The pretarsus in Aradidae (Heteroptera). — Acta Zool. Hung. 32 (3-4): 377-383.

VÁSÁRHELYI T. (1987): On the relationships of the eight aradid subfamilies (Heteroptera). — Ac- ta Zool. Hung. 33 (1-2): 263-267.

WARDLE P. (1991): Vegetation of New Zealand. — Cambridge University Press, U.K.: 1-672.

WHITE F.B. (1876): Descriptions of three new species of Hemiptera-Heteroptera from New Zealand. — Entomol. Mon. Mag. 13: 105-106.

Addresses of the Authors: Dr. Marie-Claude LARIVIÈRE Landcare Research Private Bag 92170 Auckland New Zealand E-Mail: [email protected] Mr. André LAROCHELLE Research Associate Landcare Research Private Bag 92170 Auckland New Zealand E-Mail: [email protected]

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