The Halticini of the world (Insecta: Heteroptera: Miridae: Orthotylinae ...

Zoological Journal of the Linnean Society, 2012, 164, 558–658. With 58 figures

The Halticini of the world (Insecta: Heteroptera: Miridae: Orthotylinae): generic reclassification, phylogeny, and host plant associations NIKOLAI J. TATARNIC* and GERASIMOS CASSIS Evolution & Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney 2052, Australia Received 19 January 2011; revised 2 July 2011; accepted for publication 4 July 2011

The plant bug tribe Halticini (Heteroptera: Miridae) is revised. The genus Coridromius Signoret, 1862 is removed from the Halticini to form the new monogeneric tribe Coridromiini. A generic reclassification of the Halticini is given, comprising 25 genera: Acratheus Distant, 1910; Anapus Stål, 1858; Barbarosia Kiyak, 1995; Chorosomella Horváth, 1906; Compositocoris Schwartz, Schuh & Tatarnic, 2008; Dampierella Tatarnic, 2009; Dasyscytus Fieber, 1864; Dicyphopsis Poppius, 1914; Dimorphocoris Reuter, 1890; Ectmetopterus Reuter, 1906; Euryopicoris Reuter, 1875; Goodeniaphila Tatarnic, 2009; Halticus Hahn, 1833; Labops Burmeister, 1835; Microtechnites Berg, 1883; Myrmecophyes Fieber, 1870; Nanniella Reuter, 1904; Namaquacapsus Schuh, 1974; Orthocephalus Fieber, 1858; Pachytomella Costa, 1890; Piezocranum Horváth, 1877; Plagiotylus Scott, 1874; Schoenocoris Reuter, 1890; Scirtetellus Reuter, 1890, and Strongylocoris Blanchard, 1840. Microtechnites is reinstated from synonymy with Halticus, and Platyporus Reuter, 1890 is proposed as a new junior synonym of Anapus. The following new combinations are proposed for ten species formerly placed in Halticus: Ectmetopterus bicoloratus (Kulik, 1965), Ectmetopterus comitans (Josifov & Kerzhner, 1972), Ectmetopterus fuscous (Zou, 1985), Ectmetopterus maculipes (Zou, 1985), Ectmetopterus niger (Zou, 1985) and Microtechnites bractatus (Say, 1832), Microtechnites canus (Distant, 1893), Microtechnites chrysolepis (Kirkaldy, 1904), Microtechnites minutus (Reuter, 1885), and Microtechnites spegazzini (Berg, 1883). New combinations for two species formerly placed in Cafayatina Carvalho & Carpintero, 1986 are also proposed: Microtechnites altigena Carvalho & Carpintero, 1986 and Microtechnites inesalti (Carvalho & Carpintero, 1986). Revised diagnoses are provided for each genus. Redescriptions are provided for all but the most recently described genera. A key to the genera of Halticini is provided. Colour habitus photos of the type species for most genera are given. Scanning electron micrographs of salient characters and illustrations of male and female genitalia are provided for exemplars of each of the genera. Based on our phylogenetic analysis, we define a monophyletic Halticini, with the exclusion of Coridromius. Within the Halticini, Halticus is found to be sister to the remainder of the tribe. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 558–658. doi: 10.1111/j.1096-3642.2011.00770.x

ADDITIONAL KEYWORDS: planetary biodiversity inventory – plant bugs – taxonomy.

INTRODUCTION The Halticini (Heteroptera: Miridae: Orthotylinae) was first recognized by Costa (1853) as a suprageneric group of plant bugs. Halticines are generally stout,

*Corresponding author. E-mail: [email protected]

558

dark insects, often with saltatorial hindlegs and sometimes with ant-like habitus. Many species exhibit wing polymorphism, with wing reduction in one or both sexes. All species are thought to be phytophagous, with many found on grasses (Schuh & Slater, 1995). They are mostly distributed in the Palaearctic zoogeographical region, and are particularly species-rich in the Mediterranean as well as

© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 558–658

THE HALTICINI OF THE WORLD Eastern Europe and northern Asia (e.g. Mongolia, Asiatic Russia). Most genera exhibit restricted distributions, perhaps reflecting poor dispersal ability because of the high degree of brachyptery in one or both sexes, which may be attributable to environmental stability (Southwood, 1962; Sweet, 1964; Schuh, 1974). The taxonomic position of the Halticini within the Orthotylinae has been disputed by several authors. Halticini was defined by Kirkaldy (1902) as a tribe of Orthotylinae, within which he also included the Phylinae: Pilophorini (as Pilophorinae sensu Reuter). Wagner (1973) subsequently elevated halticines to subfamilial status, segregating them on the basis of male genital structure, thereby recognizing three discrete suprageneric taxa (Orthotylinae sensu stricto, Pilophorinae, and Halticinae). Wagner (1973) further divided the Halticinae into two tribes, the nominotypical Halticini and the Chorosomellini, using characters of the head and first antennal segment. Schuh (1974) restored the Halticini as a tribe within the Orthotylinae, based primarily on the convergent lamellate parempodia. Past polemics have been recently revisited, with Kiyak (1995) restoring them to the subfamilial level. Nonetheless, most authors continue to maintain the Halticini as a tribe within the Orthotylinae (e.g. Cassis & Gross, 1995; Kerzhner & Josifov, 1999). The aim of this work is threefold: (1) ascertain if and how the Halticini are a monophyletic group; (2) determine what taxon relationships a putatively monophyletic Halticini have; and (3) establish a generic classification for the Halticini, inclusive of diagnoses and descriptions.

TAXONOMIC

559

veins not forming cells; lower portion of the head below eyes greater than twice the height of eye. This subtribe is monogeneric, only including Myrmecophyes comprised of 29 species. 2. Halticaria: small, oval, and mostly black; generally with both thin, simple setae and pale, scale-like setae; hemelytra sometimes truncate, in macropterous females barely covering abdomen, membrane when present containing two cells; head short and strongly angled backward below the midpoint, eyes overlapping anterolateral margins of pronotum; antennae long and thin, at least as long as and often longer than body, with antennal segment IV longer than entire hind leg; metafemora generally incrassate; abdomen short and rounded, without a narrowly constricted base; right paramere spoon-shaped with long base and a well-defined apical swelling (= apophysis); left paramere smaller than right, apically slender, angular, with apophysis curved down; phallotheca often black. The Halticaria includes two genera, Halticus and Barbarosia. 3. Laboparia: small to medium-sized, either macropterous and elongate-oval or brachypterous and oval, often strongly sexually dimorphic with male macropterous and female either brachypterous or macropterous; membrane with two cells; body with both simple setae and scale-like setae; head short, angled back ventrally; eyes substylate or stylate; antennae with segment III equal to or longer than segment IV; abdomen without narrow, ant-like constriction; legs often short, metafemora sometimes incrassate; right paramere spoon-shaped, longer than left, usually with apical apophysis; left paramere thin and hooked, generally with sensory lobe above base of apophysis.

HISTORY

Costa (1853) diagnosed the halticines on the basis of the tall genae, wide vertex, and saltatorial hindlegs. Schuh (1974, 1975, 1976) provided the first phylogenetic classification of the Miridae, recognizing three tribes in the Orthotylinae: Halticini, Nichomachini, and Orthotylini, all united by the possession of lyreshaped, apically convergent parempodia and male genitalia with a membranous vesica (= endosoma of Kerzhner & Konstantinov, 1999), with or without sclerotized spicules. Wagner (1973) subdivided the Halticini into three subtribes: (1) Myrmecophyaria Reuter, 1891, (2) Halticaria Kirkaldy, 1902, and (3) Laboparia Reuter, 1883, with the following diagnoses: 1. Myrmecophyaria; brachypterous morphs ant-like with a basally constricted and globular abdomen and very short hemelytra; macropterous morphs without discrete cuneus, hemelytral membrane

In this classification, the Laboparia are the largest subtribe in the Halticini, and encompass all remaining genera. Schuh (1974, 1975, 1976, 1995) did not accept this subtribal arrangement, and none of the genera described since Wagner (1973) have been separated into subtribes. Since Costa (1853), genitalic characters have played an increasing role in taxonomic delineation in the Miridae (e.g. Reuter, 1883; Knight, 1923; Wagner, 1940; Southwood, 1953; Kelton, 1955, 1959; Tatarnic & Cassis, 2008). In his seminal work on male genitalia, Kelton (1959) examined nine species from five genera of Halticini. He noted that in these taxa, the rim of the secondary gonopore is typically oval or horseshoe-shaped and the ductus seminis is a flexible cylindrical tube, both characters shared with the Orthotylini. He also observed that in the Halticini both parameres are of similar size and shape, and that except for Halticus (in his work Kelton did not


560

N. J. TATARNIC and G. CASSIS

examine any of the Halticus species that are transferred herein to Microtechnites and Ectmetopterus), the endosoma possesses several spicules. The importance of female genitalia in taxonomic delineation was first recognized by Slater (1950), who examined four genera of Halticini (Halticus, Labops, Orthocephalus, and Strongylocoris). Along with identifying the sclerotized rings of the dorsal labiate plate and structures associated with the posterior wall as useful taxonomic characters, he noted considerable variation across taxa examined, leading him to question the validity of contemporary subfamily and tribal limits. Kullenberg (1947), Kelton (1959), and Slater (1950) all had concerns regarding a classification based on pretarsal morphology, believing pretarsal structures too labile for establishing higher level taxonomic relationships. Based on their genitalic studies, Kelton and Slater both doubted the monophyly of the Halticini. Slater considered the female genitalia to be a more reliable basis for mirid classification, but for the Halticini found the structure of the posterior wall perplexing. Similarly, Kelton believed that male genitalia provided a stable taxonomic character system for the Miridae, but reported that within each tribe of the Orthotylinae remarkably divergent male genitalia were found, implying non-natural groupings. Despite these criticisms, Schuh (1974) countered that along with pretarsal form, characters of the head and the right paramere are sufficient to group the Halticini, and suggested that the posterior wall in this tribe may be considered an ‘inherently variable structure’, which eventually gave rise to the more derived form exhibited in other Orthotylinae. With respect to male genitalic structures, he argued that although the parameres may be of some use in tribal delineations, in general the male genitalia do not reveal distinctive trends within the Orthotylinae. The first formal tribal description of the Halticini to include both male and female genitalic characters was that of Schuh (1974), which included the following characters: primarily black or dark colouring, sometimes with paler markings; body robust to elongate; dorsum generally smooth and impunctate; head dorsoventrally elongate, with height of gena greater than the height of the eye; the pronotum and scutellum not greatly modified except in Myrmecophyes; commonly brachypterous in one or both sexes; metafemur frequently incrassate; vesica (= endosoma) membranous without spicules; left paramere typically elongate and apically hooked; right paramere flattened and apically expanded, generally appearing club- or spoon-shaped; female posterior wall without inter-ramal lobes; sclerotized rings variable; other

features characteristic of the Orthotylinae (e.g. lamellate parempodium). The definition of the Halticini has remained unchanged in the nearly 40 years since Schuh (1974), with many character states – in particular those of the female genitalia – unconfirmed for most genera. Additionally, more recent studies indicate another possible synapomorphy for the Halticini associated with the female vestibulum. When examining the female genitalia of Myrmecophyes oregonensis Schuh & Lattin (Schuh & Lattin, 1980), the authors observed that ‘the small sclerites which lie at the base of the anterior ovipositor valves [= first gonapophysis] in the vulvar area ventral of the bursa copulatrix are symmetrical’, in marked contrast to the asymmetrical condition expressed in the remainder of the Orthotylinae (Schuh & Lattin, 1980). In their examinations of the Halticini species Halticus luteicollis, Pachytomella parallela, and an unidentified species of Dimorphocoris, PluotSigwalt & Matocq (2006) also found the sclerites associated with the opening to the vestibulum to be symmetrical, and suggested that this might be the case for all Halticini. As we show in this work such symmetry is present in virtually all Halticini with the exclusion of Goodeniaphila, wherein the left margin of the vestibular opening has a small accessory sclerite (Tatarnic, 2009). Symmetry of the vestibular region has also been recently described in other Orthotylini, including the Hadronema group and at least one species of Araucanocoris Carvalho, 1983 (Forero, 2008), Mecomma Fieber, 1858, and Harveycapsus dimorpha Cassis, Symonds & Tatarnic (Cassis et al., 2010) (Orthotylini), suggesting that this condition may prove to be more common than first thought.

MATERIAL AND METHODS SPECIMENS EXAMINED This study was based on specimens from existing museum collections and new material collected during the course of the Planetary Biodiversity Inventory study of plant bugs (http://research.amnh. org/pbi/index.html). All efforts were made to examine the type species of all genera, and as many species within each genus as possible. In the ‘Included species’ section of each genus treatment, taxa marked with an asterisk are those that have been examined in detail. Diagnoses are provided for all genera of Halticini, whereas redescriptions are included for all genera except for those most recently described (Compositocoris, Dampierella, and Goodeniaphila). Specimens were examined from the following collections: AM, Australian Museum, Sydney; AMNH, American Museum of Natural History, New York;


THE HALTICINI OF THE WORLD ANIC, Australian National Insect Collection, CSIRO, Canberra; BMNH, Natural History Museum, London; BPBM, Bishop Museum, Honolulu; CNC, Canadian National Collection of Insects, Agriculture and AgriFood Canada, Ottawa; NHMW, Naturhistorisches Museum, Vienna; NKUM, Nankai University, Tianjin; QM, Queensland Museum, Brisbane; SAMA, South Australian Museum, Adelaide; SANC, Plant Protection Research Institute, Pretoria; TAMU, Texas A&M University, Department of Entomology, College Station; TYCN, Tomohide Yasunaga Collection, Nagasaki; ULB, Université Libre de Bruxelles, Brussels; USNM, United States National Museum of Natural History, Smithsonian Institution, Washington; ZMUC, Danish Natural History Museum, Copenhagen; ZISP, Zoological Institute, Russian Academy of Sciences, St. Petersburg.

DISSECTIONS

AND MORPHOLOGICAL ANALYSIS

Genitalia of both sexes were examined by macerating the abdomen in 5% KOH at room temperature for 5–10 min. Abdomens were then rinsed in water for approximately 10 min before being transferred to glycerol for examination, illustration, and storage. For observations of the male aedeagus it was sometimes necessary to remove the phallotheca so that fine structures of the endosoma could be documented, particularly for the secondary gonopore and spicules. As the endosoma is attached to the mesial wall of the phallotheca, it was usually necessary to tear away the latter. In other cases, the phallotheca was sufficiently translucent to observe all structures without further dissection. Dissection of females involved the removal of ventral abdominal sclerites using forceps and a minuten pin. Subsequently, a minuten was then dragged between the rami in order to separate the posterior wall from the labiate plates. Genitalic illustrations were created using a camera lucida affixed to a Leica DM LB compound microscope, at 10, 20, and 40¥ power, with 10¥ ocular lenses. Illustrations of parameres are scaled down 50% smaller than aedeagi for ease of viewing. Gross morphology was observed using a Leica MZ12 stereomicroscope. Dorsal habitus photographs were taken using the Visionary Digital BK Plus Lab photographic system (http://www.visionarydigital. com) equipped with a Canon EOS 40D camera. Because of the small size of many of these insects, multiple shots at different focal depths were taken and concatenated using HELICON FOCUS software (http://www.heliconsoft.com), and further rendered in Adobe Photoshop CS3. Scanning electron micrographs (SEMs) of gold coated and uncoated specimens were taken with a Cambridge scanning electron microscope.

561

TERMINOLOGY The terminology of morphological structures follows modern works such as Schuh (1974) and Cassis & Moulds (2002). Terminology for the genitalic structures of both sexes has varied greatly throughout the literature, leading at times to confusion in homology and terminology. Kerzhner & Konstantinov (1999) provided the most recent survey of male genitalic structures, and tabulated structural synonymies. We have taken a more conservative view of the terminology of the aedeagus, and do not follow their subdivision of the endosoma into a proximal conjunctiva and distal vesica. We therefore refer to the component of the aedeagus bounding the secondary gonopore as the endosoma, based on doubts about the homology of the vesica of these authors, as proposed by Cassis, Schwartz & Moulds (2003) and Cassis (2008). Additionally, the densely packed sclerotized rings lending structural support to the ductus seminis are herein referred to as ‘flexible ribbing’, in order to avoid confusion with the sclerotized rings of the dorsal labiate plate of the female genitalia. Information regarding the female genitalia of the Miridae is based on the comparative studies of Slater (1950) and Davis (1955). The terminology we have used is mostly derived from Davis, but with modification from Cassis et al. (2003) for the posterior wall of the bursa copulatrix, where the K-structures of Slater (1950) are referred to as the inter-ramal lobes. These are paired, petiolate, and often spinose sclerotized lobes, which project into the anterior portion of the bursa copulatrix from the dorsolateral margins of the inter-ramal sclerites. During the course of this study we identified similar inter-ramal structures in several genera of Halticini. In some cases these appear as swollen tumescences, whereas in others they are more akin to the structures characteristic of the Orthotylini, although never as narrowly basally constricted. In this study these are treated as interramal lobes, although lacking the basally constricted state expressed in the Orthotylini. In halticines, the sclerotized rings of the dorsal labiate plate (DLP) are often laterally upturned along with the adjacent margin of the DLP. The DLP also sometimes includes medial sclerotized structures (e.g. the inter-ramal bridge in Labops: Slater, 1954). In some genera the mesal margins and the rami combine with the lateral-most portion of the dorsal labiate plate to form paired medially projecting processes (MP, e.g. Anapus americanus, Fig. 10A; Labops, Figs 34G, 35A, 36E; Scirtetellus, Fig. 56D). The ventral labiate plate (VLP) is also variable, and is frequently divided, often forming a weakly sclerotized band running along the inner margins of the rami, and medially sometimes in the form of a weakly sclerotized plate. Occasionally the


562


DLP and VLP are closely adpressed in parts, and sometimes together form sclerotized folds or projections (e.g. Strongylocoris). For convenience, the external efferent system of the metathoracic scent gland is abbreviated as MTG, the first labial segment is referred to as LI, and antennal segments are abbreviated as AI through to AIV.

PHYLOGENETIC

ANALYSIS

Ninety-two morphological characters (six uninformative) were coded for 38 ingroup and five outgroup taxa (see Appendix 1 for the list of characters and character states and Appendix 2 for the data matrix). Multistate characters were treated as unordered. Rather than coding all species of Halticini we have taken an examplar approach to taxon sampling, designed to include the type species of all Halticini genera and multiple species from genera where morphological heterogeneity exists (e.g. Anapus, Dimorphocoris, Halticus, Microtechnites, Myrmecophyes, Orthocephalus, and Scirtetellus). Our sampling of Dimorphocoris and Orthocephalus was limited by a lack of availability of specimens. This taxon sampling serves as a test of Halticini and halticine generic monophyly and sister-taxon relationships, as well as providing a revised generic framework. Multiple outgroup taxa were selected from the two other tribes of Orthotylinae: Lattinova jacki Cassis, an undescribed species of Australian Orthotylus, an undescribed species of Ceratocapsus, and Aoplonema princeps (Uhler) were coded from the tribe Orthotylini, whereas the Nichomachini was represented by Nichomachus minutus. Resulting trees were rooted with Orthotylus sp. Parsimony analyses were performed in TNT 1.1, Willi Hennig Society Edition (Goloboff, Farris & Nixon, 2008). All characters were treated as unordered, with both equal weighting and implied weighting (Goloboff, 1993) approaches applied to character optimization. Tree searching was conducted using the ‘traditional’ (i.e. heuristic) search method (maxtrees = 500 000, starting Wagner trees from random seed 0, 5000 replicates, branch swapping by tree bisection-reconnection, ten trees held per replication, replacement of existing trees). Implied weights analysis used the same settings, with a concavity constant of K = 3. Jackknife values were calculated in TNT (36% removal probability, with 1000 replications). Bremer support values were calculated using the bremer.run script available from the TNT wiki website (P. Goloboff: http://tnt.insectmuseum.org).

PHYLOGENETIC RESULTS Under maximum parsimony analysis with equal character weighting three trees were found [length = 424

(excluding uninformative characters), consistency index (CI) = 0.27, retention index (RI) = 0.65, excluding uninformative characters]. The low CI is indicative of the high degree of homoplasy exhibited by multiple characters in the analysis. Nonetheless, the RI of 0.65 indicates a moderately strong phylogenetic signal. The strict consensus of these trees collapsed six branches, and has a length of 453 steps (Fig. 1). Under implied weighting a single tree was generated [Fig. 2: length (based on equal character weights) = 448, CI = 0.260, RI = 0.641]. This tree differed somewhat from those found under equal weighting, although the overall monophyly of the Halticini and most subclades were retained. Again, support values for most nodes were weak. Subtribal divisions proposed by Wagner (1973) were not supported. In their combined morphological and molecular analysis of the Cimicomorpha, Schuh, Weirauch & Wheeler (2009) presented a paraphyletic Halticini, with Halticus and Coridromius forming a sister taxon relationship separate from Compositocoris senecionus and Orthocephalus sp., the only other Halticini included in their analysis. In contrast, our results show a monophyletic Halticini, to the exclusion of Coridromius, which is found instead to be sister to Nichomachus (not included in Schuh et al., 2009). Schuh et al. (2009) provide two homoplastic morphological characters – the presence of fleshy apically convergent parempodia and the female posterior wall lacking K-structures (inter-ramal lobes) – to support a Coridromius + Halticus relationship. Both these characters are ubiquitous across the Halticini as well as some outgroup taxa (although see below our discussion of K-structures in Anapus, Labops, and Scirtetellus), and are therefore not informative in this context. Based on our phylogenetic results, we propose a monophyletic Halticini with Coridromius removed to form the monogeneric tribe Coridromiini, which is sister to the Nichomachini. Below we discuss the major findings of our phylogenetic analyses using the implied weighting phylogenetic reconstruction (Fig. 2). As we mapped only those characters whose reconstruction was unambiguous, some salient traits are not shown on the figure. This ambiguity is in many cases attributable to our treatment of multistate characters as unordered, rather than reflective of poor phylogenetic signal for the traits in question. Thus several important characters, although not appearing on Figure 2, are also discussed.

NODE A: NICHOMACHUS + CORIDROMIUS + HALTICINI Nichomachus, Coridromius, and the Halticini are united by four unambiguous synapomorphies, three of which represent aspects of the female genitalia: the


THE HALTICINI OF THE WORLD

563

Figure 1. Strict consensus of three equally parsimonious trees under equal weighting. Numbers above branches are jackknife support (36% removal probability, values below 50% not shown); numbers below are Bremer support values.

posterior wall lacks fields of minute sclerotized spines (Appendix 1: character 86-1), although these are present in some taxa; the posterior wall lacks interramal lobes (88-1), although intriguingly these are present in Anapus, Labops, and Scirtetellus; and the posterior wall lacks a medial region distinctly separated from the two lateral regions (90-1). Addi-

tionally, the lateral margins of the vulvar region (first gonapophyses) are symmetrical (91-1), although in our analysis this character is also shared with the outgroup species Aoplonema princeps. The lack of inter-ramal lobes has previously been used to define the Halticini; however here we show inter-ramal lobes to be present in three halticine genera, and a lack of


564


Figure 2. Implied weighting phylogeny (k = 3). Unambiguous character optimizations are indicated by circles: filled circles indicate synapomorphies, white circles indicate homoplasies. Numbers above and below circles indicate character number and state, respectively. Large red numbers above branches indicate jackknife support (values below 50% not reported).


THE HALTICINI OF THE WORLD these structures also occurs in Nichomachus and Coridromius; thus their absence is not exclusive to the tribe. One nongenitalic character, incrassate hind femora in females (40-1), is also synapomorphic for the group.

NODE B: NICHOMACHUS + CORIDROMIUS Nichomachus and Coridromius are united by two male genital characters: the ductus seminis is extremely short and lacks convolutions (58-0) and lacks sclerotized ribbing (59-1). Both of these traits are seen in other taxa included in this treatment. A third male genital character also helps to define this group, although in our reconstruction its reconstruction is ambiguous: only in these two taxa do we see the extreme reduction of the right paramere, which is significantly shorter than the left (50-0). In the remaining outgroup taxa the parameres are subequal in length (50-1), whereas across the Halticini the parameres are either subequal or the right is much longer than the left (50-2). Paramere size asymmetry is well known in other groups of Miridae (e.g. Dicyphina – Cassis, 1986; Termatophylini – Cassis, 1995), but is less apparent in the Orthotylinae.

NODE C: HALTICINI Characters previously used to diagnose the Halticini – the black coloration of the body (1-1), the saltatorial hindlegs (41-1), the apically clubbed or spoon-shaped right paramere (49-1), the apically hooked left paramere (54-1, 2), and the vertically elongate head (coded in our analysis as genal length, 13-2, 3) are generally true for the Halticini, although only the last of these is synapomorphic for the tribe. In the present analysis the rest of these characters exhibit high degrees of homoplasy (< 0.35 CI). Halticines are also often recognized by their robust body, although this character is diagnostic for many taxa and clearly exhibits no phylogenetic signal relative to other Miridae. Only one other unambiguous character is identified for this node: in macropterous individuals, the scutellum is not visible from above (31-0). However, visibility of the mesoscutum is variable above and below this node, and we do not consider it a particularly informative or meaningful trait. Based on limited sampling Schuh (1974: table 3) proposed that the Halticini do not possess endosomal spicules. Here we show that endosomal spicules are actually present in most halticines (72-1), although occasionally absent (spicules are lacking in Halticus, Namaquacapsus, Piezocranum and Plagiotylus). When present the endosomal spicules of the Halticini are positioned distad to the secondary gonopore (73-

565

1), in contrast to the Orthotylini, in which they are attached basad to the gonopore. Symmetry of the margins of the vestibular opening (first gonapophyses: 91-1) has recently been proposed as a potential synapomorphy for the Halticini (PluotSigwalt & Matocq, 2006). However, although common to all Halticini, this condition is also exhibited in several outgroup taxa included in our analysis – Coridromius, Nichomachus, and Aoplonema princeps (as well as the rest of the Hadronema group: see Forero, 2008) – and cannot therefore be considered synapomorphic for the Halticini.

TAXONOMY CORIDROMIINI

NEW TRIBE

Type genus: Coridromius Montrouzier. Type species: Ocypus variegatus Montrouzier, 1861 (monotypy). Ocypus Montrouzier, 1861: 67, junior homonym of Ocypus Leach, 1819 (Coleoptera). Coridromius Signoret, 1862: 5 (nom. nov. for Ocypus Montrouzier, 1861). Neocypus Distant, 1914: 378 (unnecessary nom. nov. for Ocypus Montrouzier, 1861). Carvalho, 1987: 61; Linnavuori, 1994: 15; Schuh, 1995: 46; Cassis & Gross, 1995: 185 [Australian catalogue (cat.); transferred to Halticini]; Schuh, 1995: 46 (world cat.; Halticini); Liu & Zhao, 1999: 55; Miyamoto & Yasunaga, 1999: 33 (descr.); Chérot, Konstantinov & Yasunaga, 2004: 57 (descr.). Coridromoides Carvalho, 1956: 54 (gen. nov.; type species: Coridromoides carinatus Carvalho, 1956, by original designation); Tatarnic & Cassis, 2008: 1 (descr.; generic revision). Diagnosis: Readily distinguished from all other Miridae by the following characters: male with hypodermic genitalia formed by the coupling of the left paramere and the aedeagus; female genitalia extremely reduced, with subgenital plate absent, vulvar region unsclerotized and symmetrical, posterior wall simple and lacking medial area and interramal lobes. Additionally, body short and squat, metafemora greatly swollen and modified for jumping, hemelytra strongly deflected at cuneus. Diversity and distribution: Coridromius currently includes 33 species found throughout the Old World tropics and subtropics, with the Australian species Coridromius chenopoderis Tatarnic & Cassis introduced to Hawaii, south-western USA, and northern Mexico (Tatarnic & Cassis, 2008).


566


Remarks: In our analysis Coridromius does not fall within the Halticini. This is unsurprising, as it lacks many of the characteristics typical of the tribe: elongate bases to both parameres, a spoon-shaped right paramere, mostly black coloration, the aedeagus with a well-developed endosoma (but see Halticus), the ductus seminis with sclerotized ribbing and sclerotized secondary gonopore, and well-developed sclerotized rings with upturned lateral margins. Indeed, most genitalic characters in Coridromius are unlike those of any Halticini (or any other mirid): the female genitalia are highly reduced with the sclerotized rings completely absent, and in the male the aedeagus and left paramere are coupled to form a piercing intromittent organ. Coridromius is the only plant bug genus known to practice traumatic insemination, whereby males use hypodermic genitalia to stab females in the abdomen during mating, completely bypassing the female genital tract (Tatarnic et al., 2006). As a result, females have extremely reduced genitalia, with some species exhibiting various paragenital modifications at the site of insemination, believed to reduce matinginduced costs (Tatarnic et al., 2006, Tatarnic & Cassis 2010). Because of this, the use of genitalic traits – in particular genital reduction or simplification – should not be taken as evidence of common ancestry. We believe that Coridromius is sufficiently different from all other Halticini (as well as all other existing tribes within the Orthotylinae) to justify the erection of its own monotypic tribe. Ultimately its relationship to other tribes of Orthotylinae should not be determined through morphology alone, but will instead necessitate inclusion of other data (i.e. DNA). A thorough revision of the genus can be found in Tatarnic & Cassis (2008).

HALTICINI (Numbers following generic names indicate numbers of included species.)

CHECKLIST

OF GENERA OF

HALTICINI

Acratheus Distant, 1910 [3 species] Oriental and African region Anapus Stål, 1858 = Platyporus Reuter, 1890; new synonymy [8] Holarctic Barbarosia Kiyak, 1995 [2] Palaearctic region Chorosomella Horváth, 1906 [2] Palaearctic region Compositocoris Schwartz, Schuh & Tatarnic [1] Ethiopian Region Dampierella Tatarnic, 2009 [1] Australia Dasyscytus Fieber, 1864 [1] Palaearctic region

Dicyphopsis Poppius, 1914 [4] Ethiopian region Dimorphocoris Reuter, 1890 [52] Palaearctic and African region Ectmetopterus Reuter, 1906 [6] Palaearctic and Oriental region Euryopicoris Reuter, 1875b [2] Palaearctic region Goodeniaphila Tatarnic [2] Australia Halticus Hahn, 1832 [18] Eastern Hemisphere and North America Labops Burmeister, 1835 [12] Holarctic Microtechnites Berg, 1879 [5] reinstated status Neotropical and Nearctic regions Myrmecophyes Fieber, 1870 [27] Holarctic Namaquacapsus Schuh, 1974 [1] Ethiopian region Nanniella Reuter, 1904 [6] Ethiopian region Orthocephalus Fieber, 1858 [23] Palaearctic and Nearctic regions Pachytomella Reuter, 1890 [7] Palaearctic region Piezocranum Horváth, 1877 [4] Palaearctic region Plagiotylus Scott, 1874 [7] Palaearctic region Schoenocoris Costa, 1842 [1] Palaearctic region Scirtetellus Reuter, 1890 [14] Palaearctic region Strongylocoris Blanchard, 1840 [16] Palaearctic region

HALTICINI COSTA (FIGS 3–58) Halticini Costa, 1853: 75 (new tribe); Kirkaldy, 1906: 130 (cat.); Knight, 1923: 497, 498 (gen. key); Blatchley, 1926: 797 (east US gen. key); Hsiao, 1942: 253 (Chinese gen. key); Wagner, 1952: 95 (desc.; Palaearctic gen. key); Carvalho & Leston, 1952: 245 (British gen. key); Carvalho, 1952: 39, 40, 73 (list); Carvalho, 1955: 65 (world gen. key); Carvalho, 1958: 5 (cat.); Slater, 1950: 43 (female genitalia); Kelton, 1959: 33 (male genitalia); Wagner, 1961: 47 (European fauna); Wagner & Weber, 1964: 250 (French fauna); Wagner, 1973: 2 (Mediterranean fauna); Schuh, 1974: 26, 273 (classification); Schuh, 1975: 13 (femoral trichobothria); Schuh, 1976: 18, 35 (pretarsus; classification); Linnavuori, 1994: 4 (West African gen. key); Schuh & Slater, 1995: 44 (world cat.); Kerzhner, 1988: 778, 826 (key to gen.; Palaearctic fauna); Kerzhner & Konstantinov, 1999: 124 (male genitalia). Halticocoridae Douglas & Scott, 1865: 35 (fam. stat.). Stiphrosomatidae Douglas & Scott, 1865: 35 (new fam.). Halticaria Kirkaldy, 1902: 139 (list); Reuter, 1910: 115 (desc.; key); Poppius, 1911: 33 (class.); Poppius, 1914: 59, 82 (key; descr.); Van Duzee, 1916: 43 (cat.); Butler, 1923: 463 (descr.; key); Wagner, 1973: 15 (European fauna). Diplacaria Reuter, 1883: 564: (new division). Laboparia Reuter, 1883: 567 (new division; gen. key); Puton, 1886: 53 (cat.); Atkinson, 1890: 117 (cat.);



KEY 1. – 2. – 3.

– 4.

– 5.

–

6. – 7. – 8. – 9. – 10. – 11. – 12. – 13. – 14. –

TO THE GENERA OF

567

HALTICINI

Scutellum, pronotum, hemelytron, and thoracic pleura densely punctate (Figs 5A–C, E–F, 43A–F); ductus seminis appears to open into sclerotized tube running to apex of endosoma (Figs 6A, 44A–B)..................................2 Scutellum, pronotum, hemelytron, and thoracic pleura not all densely punctate; male secondary gonopore variable.........................................................................................................................................3 Frons and vertex deeply punctate; left paramere broad, almost as broad as right (Fig. 6B, C)...........Acratheus Frons and vertex not deeply punctate; left paramere with twisted apophysis, unlike right paramere (Fig. 44C– E) ..................................................................................................................................... Nanniella Both sexes strongly ant-mimetic, sometimes with clear ant-like waist (Fig. 4: Myrmecophyes alboornatus). Macropterous individuals without distinct cuneus, membrane without cells. Height of gena greater than twice height of eye (Fig. 39C)...................................................................................................Myrmecophyes Ant-mimetic or not, but never with an ant-like waist; macropterous individuals with cuneus, membrane with one or two enclosed cells; height of gena variable ....................................................................................... 4 Pale; both sexes large, long, and slender (Fig. 3: Chorosomella jakowleffi); hemelytra in macropterous male not covering apex of abdomen, hemelytra of female reduced and pad-like; antennae long, with AI longer than head and pronotum combined; vertex level with prominent conical projection (Fig. 13A, B) ................. Chorosomella Usually dark; sometimes elongate; in macropterous individuals apex of abdomen always covered; antennae variable but AI not longer than head and pronotum combined; vertex never with conical projection .............. 5 Eyes stylate and angled upwards (Fig. 33A–C); head tall (Fig. 33B, C); male endosoma with tightly packed mass of thin, needle-like spicules (Figs 34A, 36B); female posterior wall with inter-ramal lobes (Figs 34H, 35B, 36G); DLP sometimes with inter-ramal bridge (Figs 34G, 36E, F); VLP + rami sometimes forming medially projecting sclerotized lobes (Figs 34G, 35A, 36E) ......................................................................................... Labops Eyes not stylate and angled upwards; head tall or short; male endosoma without tightly packed mass of needle-like spicules; female genitalia with or without inter-ramal lobes on posterior wall, inter-ramal bridge, or medially projecting sclerites formed by VLP and rami ........................................................................... 6 Small and gracile; both sexes macropterous, with hemelytra semitranslucent; eyes round and bulging (Fig. 20A– D); pronotum campanulate with lateral margins distinctly concave (Fig. 20A)..............................Dicyphopsis Size variable, robust; degree of wing reduction variable, hemelytra always opaque; eyes variable; pronotum variable.........................................................................................................................................7 Posterior margin of head closely appressed to pronotum, with posterior of eyes touching or nearly touching anterior of pronotum (Figs 28A, 47B, 57A); MTG sometimes obsolete (e.g. Figs 47D, E, 57C, D)....................8 Posterior margin of head straight or convex, not concave, with posterior of eyes not touching anterior of pronotum; MTG external efferent system always well developed...........................................................................10 Dimorphic, macropterous male narrow-bodied with parallel-sided hemelytra, brachypterous female pear-shaped (Fig. 4: Pachytomella passerini); pronotum (Fig. 47B) and hemelytra impunctate ........................ Pachytomella Both sexes macropterous and broadly oval; pronotum and hemelytra punctate (Figs 17A, 57A).....................9 MTG obsolete (Fig. 57C, D); margins of opening to female vestibulum (first gonapophyses) symmetrical (Fig. 58F)......................................................................................................................Strongylocoris MTG well developed (Fig. 28D); opening to female vestibulum with small accessory sclerite on left margin (Fig. 29G).....................................................................................................................Goodeniaphila Posterior margin of vertex carinate (Figs 17A, 24A, 30A, 37A, 45A–C, 50B) ............................................. 11 Posterior margin of vertex not carinate (Figs 7A, 11A, 15A–C, 22A, 26B, 39A, 52B, 54A) ........................... 16 Head in macropterous individuals with prominent transverse sulcus across frons and vertex, much less distinct in brachypterous individuals (Fig. 4: Piezocranum simulans)....................................................Piezocranum Head without transverse sulcus spanning vertex and frons...................................................................12 Membrane of macropterous individuals covered with dense, short setae; in macropterous female abdomen extends laterally beyond hemelytra (Fig. 3: Dasyscytus sordidus); left paramere distinctly folded (Fig. 19B)...Dasyscytus Membrane of macropterous individuals never covered with setae, abdomen of macropterous female never extending laterally beyond hemelytra; left paramere not folded ............................................................. 13 Antennal insertion in front and above lower margin of eye (Figs 24B, 30B, D, 37B–C); antennae usually longer than body (at least in macropterous individuals).................................................................................14 Antennal insertion in line with or well below lower margin of eye (Figs 17B, C, 41A, 45B, C, 50B); antennae always shorter than body................................................................................................................22 Aedeagus with scoop-shaped secondary gonopore and extremely reduced endosoma without spicules (Fig. 31B, C); sclerotized rings of DLP large and subcontiguous (Figs 31F, 32A)...................................................Halticus Aedeagus without distinctly scoop-shaped secondary gonopore, endosoma well developed, with or without spicules; sclerotized rings moderately sized, widely separated ............................................................................ 15


568


15. Macropterous; hemelytra without pale, scale-like setae; right paramere projecting out of pygophore (Fig. 24G, H); pygophore margin strongly concave below left paramere (Fig. 24H); left paramere trifurcate (Fig. 25B); endosoma without several elongate spicules (Fig. 25A) ........................................................................ Ectmetopterus – Macropterous or brachypterous; macropterous hemelytra with pale, scale-like setae (Fig. 37A); right paramere not projecting from pygophore (Fig. 37G, H); pygophore margin straight, not sulcate (Fig. 37G, H); left paramere L-shaped; endosoma with several, often serrate, elongate spicules (Fig. 38A)............................Microtechnites 16. Posterior region of pronotum rugulopunctate (Figs 11A, 26B); hemelytra punctate; coleopteroid (Fig. 3: Barbarosia punctulata, Euryopicoris nitidus; Fig. 26A).........................................................................................17 – Posterior of pronotum not rugulopunctate; hemelytra not punctate; degree of wing reduction variable..........18 17. Fore- and mesotibiae thickened and distinctly curved; female sclerotized rings small and simple ... Euryopicoris – Fore- and mesotibiae not thickened and distinctly curved; female sclerotized rings complex ............ Barbarosia 18. Mostly black with some yellow or orange markings; usually with hemelytra reduced; metafemora incrassate; posterior wall of bursa copulatrix usually with paired inter-ramal lobes (Figs 8E, 9F, 10B, 55D, 56C) .......... 19 – Mostly brown or green or pink; sometimes brachypterous; metafemora not incrassate; posterior wall of bursa copulatrix without inter-ramal lobes..................................................................................................20 19. Mainly brachypterous, males rarely macropterous; sometimes clothed with pale, scale-like setae; aedeagus with elongate subapical portion of ductus seminis leading to secondary gonopore without flexible ribbing, weakly to strongly sclerotized (Figs 8A, 9A)...............................................................................................Anapus – Staphylinoid, hemelytra reduced to pads covering the base of the abdomen; ductus seminis with flexible ribbing through entire length basal to secondary gonopore (Fig. 55A).....................................................Scirtetellus 20. Mostly green and yellow, hemelytra tinted pink; male macropterous and elongate, female staphylinoid and broadly oval; female with apically stellate setae (Fig. 15A, B, F) ........................................... Compositocoris – Mostly brown with tan markings; hemelytra without pink tint; male macropterous and elongate or brachypterous, female brachypterous and pear-shaped to broadly oval; female without apically stellate setae.....................21 21. Male always macropterous, female brachypterous; MTG evaporative area broad and triangular, peritreme rounded (Fig. 52E)............................................................................................................Schoenocoris – Male brachypterous or macropterous, female brachypterous; MTG evaporative area tongue-like, peritreme a narrow strip (Fig. 22D, E)...............................................................................................Dimorphocoris 22. Mostly black or dark brown; usually with white scale-like setae (e.g. Fig. 17A–D).....................................23 – Pale green and yellow or chestnut, sometimes with red markings; without white scale-like setae ................ 24 23. Clothed in white, scale-like setae (Fig. 17A–D); both sexes macropterous; secondary gonopore scoop-shaped (Fig. 18C); gena height slightly less than eye height (Fig. 17B, C).............................................Dampierella – Often clothed in white, scale-like setae; both sexes macropterous or brachypterous; secondary gonopore basally constricted to form dorsoventral bowl shape, with apex dorsoventrally compressed and laterally expanded (Fig. 46A, B); gena height slightly greater than eye height (Fig. 45B)......................................Orthocephalus 24. Pale green, sometime with yellow and black markings; dimorphic, male macropterous, female coleopteroid; MTG obsolete (Fig. 50C, D)..........................................................................................................Plagiotylus – Predominantly chestnut-coloured with red corium at base and apex.....................................Namaquacapsus

Reuter, 1891: 17 (descr.); Distant, 1904: 479 (descr.); Reuter, 1905a: 19, 28 (descr.; key); Reuter, 1909: 71 (list); Oshanin, 1910: 793 (Palaearctic cat.); Oshanin, 1912: 73 (Palaearctic cat.); Van Duzee, 1916: 211, 373 (key; descr.); Wagner, 1973: 15 (subtribe; European fauna). Labopini Knight, 1923: 501 (key); Blatchley, 1926: 797 (east US gen. key); Hedicke, 1935: 53 (gen. key); Knight, 1941: 8, 19, 74 (Illinois gen. key); Kiritshenko, 1951: 117 (USSR gen. key). Myrmecophyaria Reuter, 1891: 106 (new division); Wagner, 1973: 15 (subtribe; European fauna). Halticarini Zimmerman, 1948: 198 (key). Diagnosis: Usually black or dark brown with or without lighter contrasting markings, sometimes

mostly light brown, tan or green; body generally stout and compact, sometimes elongate; hindlegs often saltatorial; pronotal texture most often smooth and glossy, sometimes punctate; vestiture variable, either only with thin simple setae, or also with pale and lamellate setae; legs and antennae often spinose; head typically dorsoventrally elongate, height of genae almost always equal to or greater than height of eye; pronotum campanulate, rectangulate or trapezoidal; aedeagus with thinly sclerotized phallotheca, apex of phallotheca most often constricted, sometimes keeled, rarely with thin apical projections, ductus seminis long or short with flexible ribbing formed by numerous thin, closely packed sclerotized rings, sometimes with elongate subapical segment lacking flexible ribbing; secondary gonopore weakly to



Figure 3. Photographs of Halticini genera: Acratheus–Halticus. Abbreviations: F, female; M, male.


569

570


Figure 4. Photographs of Halticini genera: Labops–Strongylocoris. Abbreviations: F, female; M, male.


THE HALTICINI OF THE WORLD strongly sclerotized, often with distinctive scale-like texturing, endosoma almost always present as a membranous bag within the phallotheca (except in Halticus), often with one or several sclerites, sometimes with fields of spines or dentitions; basal portion of both parameres usually elongate (except in Dampierella and Goodeniaphila, where the base of the left paramere is relatively short with respect to other halticines), both parameres ventrally concave; left paramere usually L-shaped, sometimes with swollen sensory lobe, with long apophysis, often apically hooked and sometimes bifid; right paramere flattened and generally spoon- or club-shaped, sometimes with a small apical apophysis; posterior wall of female without inter-ramal lobes (but see Anapus, Scirtetellus, and Labops); sclerotized rings variable, but generally weakly concave or laterally upturned; margins of first gonapophyses symmetrical, sometimes with weak sclerotization.

ACRATHEUS DISTANT (FIGS 3, 5–6) Acratheus Distant, 1910a: 16 (gen. nov. type species: Acratheus nocturnus Distant, 1910 by monotypy); Distant, 1910b: 284 (India); Reuter, 1910: 161 (cat.); Carvalho, 1952: 73 (list); Carvalho, 1955: 67 (key); Carvalho, 1958: 6 (cat.), Linnavuori, 1994: 7 [diagnosis (diag.), key to spp.]; Schuh, 1995: 45 (cat.) Diagnosis: Acratheus is distinguished from other Halticini except Nanniella by the densely punctate scutellum, pronotum, and thoracic pleura. It differs from Nanniella by the lack of deep punctures on the frons and vertex and the shape of the left paramere, which in Nanniella is more elongate and apically barbed. Redescription: Macropterous, body length 2–3 mm. Coloration (Fig. 3): mostly dark brown to black, with yellow or yellow-brown markings. Surface and vestiture (Figs 3, 5A–H): head mostly smooth with shallow punctures limited to genae (Fig. 5A–C). Pronotum, scutellum, and thoracic pleura densely punctate (Figs 3, 5A–C, E–F) Hemelytra densely punctate, less so towards apex of corium (Figs 3, 5A); cuneus punctate. Abdominal sternites with few punctures laterally (Fig. 5H). Thorax and abdomen sparsely covered with longish, brown, decumbent setae (Fig. 5B, E, H). Structure: body elongate and slightly ovate. Head (Figs 3, 5A–C): transverse, short, taller than wide; eyes large, height of gena slightly greater than eye height; posterior margin of vertex carinate, sublinear with eyes not touching pronotum; vertex convex; clypeus not distinctly projecting forward; maxillary plate long and vertical; bucculae small, narrow. Antennae (Figs 3, 5A–C): insertion in line with mid-

571

point of eye; elongate, narrow, AI only slightly thicker than AII. Labium (Fig. 5B): extending beyond metacoxae, LI somewhat swollen and elongate; thorax (Figs 3, 5B, D–F): pronotum trapezoidal, steeply inclined, collar flat, visible from above, posthumeral angles distinctly depressed, posterior margin carinate, medially concave; mesoscutum visible; scutellum medially raised, basally depressed, laterally declivent; metathoracic spiracle exposed, operculum large, without evaporative bodies; MTG external efferent system reduced, tongue-like, peritreme located along posterior margin of metepisternum, narrow strip of evaporative areas anteriad to peritreme. Hemelytra (Figs 3, 5A): Laterally declivent at corial fracture, lateral margin slightly flared apically; membrane extending beyond abdomen. Legs (Figs 3, 5G): elongate, narrow; metafemora not incrassate; pretarsi without pulvilli. Abdomen: elongate-ovoid. Male genitalia (Figs 6A–C, 5H): pygophore conical, simple, without modifications or processes; parameres subequal in length, both with elongate sensory lobe; left paramere broad, with thin, strongly arcuate distal arm; right paramere spoon-shaped with thin apical apophysis, projecting beyond genital opening of pygophore; phallotheca simple, elongate, abruptly narrowing from midpoint to apex; ductus seminis short, with irregular U-shaped sclerotized secondary gonopore, opening into a conical sclerotized tube running length of phallotheca. Female genitalia (Fig. 6D, E): sclerotized rings minute, thin, lateral margins and adjacent portion of DLP weakly upturned; posterior wall membranous, with very weak lateral swellings; margins of vestibulum symmetrical with medial margins of first gonapophyses forming slightly tumescent, weakly sclerotized plates. Diversity and distribution: Acratheus has four species, two in India and two in equatorial Africa. Lack of records despite intensive sampling in Middle Asia by Linnavuori (1975, 1984, 1986, 1992) suggests this apparent geographical disjunction is real. Included species: Acratheus albipes (Motschulsky, 1863) India Acratheus nocturnus Distant, 1910 India Acratheus ocellaris Linnavuori, 1994* Benin; Ivory Coast; Nigeria; Togo Acratheus punctiger (Linnavuori, 1975)* Benin; Nigeria; Sudan; Upper Volta Biology and host plant associations: Acratheus is found on undergrowth in savannah and rain forests (Linnavuori, 1994). No host plants have been recorded for this genus.


572


Figure 5. Scanning electron micrograph images of Acratheus ocellaris. A, dorsum; B, head and thorax, lateral view; C, head, dorsal view; D, detail of evaporative bodies; E, thoracic pleura; F, external efferent system of the metathoracic scent gland; G, tarsus; H, pygophore.

Remarks: Acratheus is one of only two halticine genera characterized by deep punctations on the head, pronotum, thoracic pleura, and hemelyra, the other being the endemic African genus Nanniella. Schuh (1974) first proposed a close relationship between these genera. In contrast, Linnavuori (1994) stressed that Nanniella ‘undoubtedly represents a

separate evolutionary lineage’, and posited that Acratheus and Halticus are closely related, based on the following shared similarities: an elongate head, swollen first labial segment (LI), and halticine-type parameres. He also noted that in Nanniella the head is notably shorter and LI is only slightly thicker than LII.



573

Figure 6. Male and female genitalia of Acratheus ocellaris. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, first gonapophyses; F, posterior wall. Abbreviations: DS, ductus seminis; PB, phallobase; PT, phallotheca; SG, secondary gonopore; SR, sclerotized ring.

Based on our observations and phylogenetic results (Figs 1, 2), we concur with Schuh (1974) that Acratheus and Nanniella are closely related, and do not see the differences suggested by Linnavuori. The head projects forward only a little more and the first labial segment is only slightly less thick in Nanniella than in Acratheus. Conversely, both genera share many characters, with the following significant similarities: the posterior margin of the pronotum is concave; the configuration of the external efferent system of the metathoracic gland; the secondary gonopore is wide and distally modified into a sclerotized tube running the length of the phallotheca; the shape of the parameres; the posterior wall of the bursa copulatrix is membranous to weakly sclerotized, bearing two minor lateral swellings; and the sclerotized rings of the female dorsal labiate plate are weakly sclerotized and upturned laterally. This redescription of Acratheus is based on examination of the African species (Ac. ocellaris and Ac. punctiger) and the literature (Linnavuori, 1994). We did not have access to the Indian species Ac. albipes and Ac. nocturnus; however, Linnavuori (1994)

described the parameres of the former species, which are very similar to the African species.

ANAPUS STÅL (FIGS 3, 7–10) Anapus Stål, 1858: 188 (gen. nov.; type species: Anapus kirshbaumi Stål, 1858: 189 by monotypy); Walker, 1873: 160 (list); Reuter, 1891: 69, 159 (key; descr.); Kirkaldy, 1906: 131 (list); Hueber, 1906: 5 (key); Oshanin, 1910: 787 (cat.); Reuter, 1910: 147 (cat.); Stichel, 1933: 235 (key); Kiritshenko, 1951: 126 (key); Wagner, 1952: 96, 106 (key; descr.); Carvalho, 1955: 66 (key); Carvalho, 1958: 6 (cat.); Kerzhner, 1964a: 965 (diag., key to spp.); Wagner, 1973: 55 (descr., key to spp.); Schuh, 1995: 45 (world cat.). Labops (Merotrichaea) Reuter, 1875c: 24 (subgen. nov.; type species: Orthocephalus freyi Fieber, 1864, by subsequent designation Kirkaldy, 1906); Reuter, 1875b: 87 (key); Reuter, 1891: 79 (synonymy). Merotrichia Carvalho, 1958: 6 (cat.; generic status). Platyporus Reuter, 1890: 246 (gen. nov.; type species: Platyporus dorsalis Reuter, 1890 by monotypy); Carvalho, 1958: 30 (cat.); Wagner, 1973: 53 (descr.); Schuh, 1995: 69 (world cat.). New synonymy.


574


Figure 7. Scanning electron micrograph images of Anapus longicornis (male). A, dorsum; B, lateral view; C, head and thorax, lateral view; D, external efferent system of the metathoracic scent gland; E, venter; F, tarsus; G, abdomen, lateral view.

Diagnosis: Recognized by the following combination of characters: substylate eyes; antennal insertion below ventral margin of eye; AI swollen and longer than height of eye, with several elongate spines; hemelytra of brachypterous individuals very short; male ductus seminis long, with elongate apical section lacking ribbing; secondary gonopore rounded, opening ven-

trally, with prised operculum; and, female posterior wall of bursa copulatrix often with bilaterally symmetrical, sclerotized inter-ramal tumescences. Redescription: Coloration (Fig. 3): Body uniformly black with minor yellow and orange markings; appendages black. Surface and vestiture (Figs 3,



575

Figure 8. Male and female genitalia of Anapus kirshbaumi. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix (dorsal view); E, posterior wall. Abbreviations: EN, endosoma; IRL, inter-ramal lobe; SG, secondary gonopore; SP, spicule.

7A–E, G): Body impunctate, sometimes pronotum, scutellum and hemelytra rugulose, often with wrinkles on vertex and frons radiating from midline. Body with erect and semi-erect simple setae, sometimes intermixed with dense decumbent, white, scalelike setae. Head with a few elongate thin erect black setae. Antennae with semi-erect, spine-like setae; AI with a few to numerous elongate spines, especially in males. Legs with elongate sharp spines, particularly

on metatibiae. Structure: both sexes usually brachypterous, rarely macropterous. Head (Figs 3, 7A–C, E): transverse, triangular in shape in dorsal view, with posterior margin straight to very weakly concave; head in lateral view angled somewhat caudally below antennae; height of head below eye approximately twice height of eye; clypeus strongly declivent, not projecting forward; posterior margin of vertex not carinate; frons rounded, strongly declivent


576


Figure 9. Male and female genitalia of Anapus dorsalis. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, bursa copulatrix, ventral view; F, posterior wall, ventral view.

in lateral view; posterior margin of head wider than anterior of pronotum; eyes substylate; Antennae (Figs 3, 7A–C, E): antennal insertion well below eye; subequal to body length; AI thickened, sometimes incrassate; remaining segments thin. Labium (Fig. 7E): length variable, reaching from mesocoxae to basal abdomen sternites; LI incrassate. Thorax (Figs 3, 7A–D): pronotum short, weakly declivent in lateral view, rectangular in brachypterous forms, trapezoidal in male macropterous individuals; collar absent; callosite region sometimes distinct, lateral

margins rounded; posterior margin weakly concave; mesoscutum only visible in macropterous individuals; scutellum triangular, flat to tumescent; metathoracic spiracle large and exposed, sometimes with evaporative bodies along posterior margin; MTG external efferent system broad, swollen and triangular, generally reaching lower margin of metathoracic spiracle; peritreme oval, medially on metepisternum, subparallel to posterior margin of pleuron. Hemelytra (Figs 3, 7A–B): usually brachypterous, sometimes reduced to a small pad, in macropterous morphs with



577

Figure 10. Female genitalia of Anapus americanus. A, bursa copulatrix, dorsal view; B, posterior wall; C, first gonapophyses, ventral view. Abbreviations: IRL, inter-ramal lobe; SR, sclerotized ring.

single membrane cell, minor cell obsolete. Legs (Figs 3, 7A–B, E–F): metafemora strongly incrassate; foretibiae usually swollen apically and weakly arcuate; pretarsi with elongate fleshy parempodia (except Anapus americanus). Abdomen (Figs 3, 7G): pear-shaped, elongate oval, or oval. Male genitalia (Figs 8A–C, 7B, E, G, 9A–C): pygophore triangular, posterior margin concave at left paramere insertion; right paramere sometimes projecting slightly from pygophore; phallotheca simple, sometimes moderately sclerotized apically; left paramere sensory lobe broad, apophysis thin, hooked; right paramere broad, spoonshaped, with small apical apophysis; phallotheca simple, without processes; ductus seminis elongate with flexible ribbing, sometimes very slender (as in Anapus freyi), subapically with elongate section, sometimes heavily sclerotized, lacking flexible ribbing; secondary gonopore elongate, U-shaped, with semiclosed operculum; endosoma with one or two simple spicules of varying size and shape, in most species also with paired, elongate, and sclerotized spiculate strips near apex. Female genitalia (Figs 8D, E, 10A–C, 10A–F): sclerotized rings moderate to large, elongate, lateral margins weakly upturned

along with adjacent portion of DLP, DLP sometimes with heavily sclerotized inter-ramal bridge; margin of VLP adjacent to rami sclerotized, medially with a small flange; posterior wall of bursa copulatrix variable, but always with bilaterally symmetrical, sclerotized structures (inter-ramal tumescences), sometimes toothed or serrate; margins of vestibular opening symmetrical with weak sclerotization. Diversity and distribution: Anapus has seven Holarctic species, one from the western USA and the remainder Palaearctic. Included species: Anapus americanus Knight, 1959* USA Anapus dorsalis (Reuter, 1890)* comb. nov. Russia; Turkey Anapus freyi (Fieber, 1864)* Russia = Pachytoma jakovloeffi Reuter, 1879 (syn. Reuter, 1881) Anapus kirschbaumi Stål, 1858* Russia = Orthocephalus opacus Jakovlev, 1875 (syn. Jakovlev, 1877)


578


= Anapus pectoralis Horváth, 1904 (syn. Kerzhner, 1996) Anapus longicornis Jakovlev, 1882* Hungary; Russia = Labops (Pachytoma) arenarius Horváth, 1884 (syn. Reuter, 1890) = Anapus flavicornis Reuter, 1904 (syn. Kerzhner, 1970) Anapus pachymerus (Reuter, 1881) Spain Anapus rugicollis (Jakovlev, 1877)* Russia = Pachytoma sibirica Sahlberg, 1878 (syn. Kerzhner, 1962) = Pachytoma nigrita Jakovlev, 1881 (syn. Kerzhner, 1962) Biology and host plant associations: Host plant records are known for four species of Anapus (Table 1; Schuh, 1995). Anapus freyi is known from a grass species, and the other three species are known from either euasterid or eurosid angiosperms. Remarks: Herein we propose Platyporus as a junior synonym of Anapus, resulting in the new combination: Anapus dorsalis (Reuter, 1890). This species conforms to the generic diagnosis of Anapus given in this work, and shares significant overlap in salient morphology, and striking similarities in the male aedeagus and female posterior wall of the bursa copulatrix. This synonymy is supported by our phylogenetic analysis. In all the Old World species of Anapus examined the male genitalia are nearly identical, with the following similarities: the distal portion of the ductus seminis is sclerotized, the endosoma is relatively small, bearing one or two spicules and apically with sclerotized fields of small spines, and the left paramere has a broad sensory lobe. However, in the New World species, An. americanus, the distal portion of the ductus seminis is weakly sclerotized, the endosoma is much larger with the apical fields of small spines only faintly visible, and the left paramere lacks the large sensory lobe present in other species. Additionally the body is much more ant-like in shape, the hemelytra are reduced to very small pads, the pretarsi lack fleshy pulvilli, and the first antennal segment is thinner, more elongate, and has fewer black spines. Despite these differences we consider An. americanus to represent a divergent New World lineage within Anapus rather than a new monotypic genus, based primarily on commonalities in the structure of the female posterior wall of the bursa copulatrix. Again, this is supported by our phylogenetic results. Anapus shares some notable female genitalic peculiarities with several other taxa: in particular, the sclerotized inter-ramal bridge of the DLP and the

inter-ramal lobes of the posterior wall (characters 82 and 88) are both shared with Labops (Slater, 1950; Slater, 1954), with the latter of these structures also found in Scirtetellus. In these other genera and An. americanus, the inter-ramal lobes are hollow, apically tapered, and medially convergent, and spinose. These appear homologous to the inter-ramal lobes found in the Orthotylini (Slater, 1950; also known as K-structures), although they differ in that they are generally broader and much less narrowly constricted basally (89-1). Aside from An. pachymerus, all species of Anapus were examined for this study.

BARBAROSIA KIYAK, 1995 (FIGS 3, 11–12) Halticidea Reuter, 1901: 172 (gen. nov.; junior homonym of Halticidea Hohrn, 1893 [Coleoptera]; type species: Halticidea punctulata Reuter, 1901 by monotypy); Kirkaldy, 1906: 130 (list); Oshanin, 1910: 802 (Palaearctic cat.); Reuter, 1910: 147 (cat.); Kiritshenko, 1951: 127 (key); Carvalho, 1952: 73 (list); Carvalho, 1955: 68 (key); Carvalho, 1958: 11 (cat.); Wagner, 1973: 13 (descr.; key); Schuh, 1995: 53 (world cat.). Barbarosia Kiyak, 1995: 216 (nom. nov. for Halticidea Reuter, 1901). Diagnosis: Barbarosia is recognized by the following combination of characters: coleopteroid; mostly shiny black; white scale-like setae on laterotergites; frons extending beyond clypeus; eyes substylate; antennae short, subequal to body length; left paramere L-shaped; endosoma with single S-shaped spicule; and, female DLP exaggerated, sclerotized ring strongly convoluted. It is similar to Halticus but with shorter antennae, which in the male are slightly longer, and in the female slightly shorter than the body. The female sclerotized rings are unlike those of any other halticine. Redescription: Coleopteroid, 2.3–3.35 mm, male slightly smaller than female. Coloration (Fig. 3): body mostly shining dark brown to black with paler markings. Surface and vestiture (Figs 3, 11A–E): body sparsely clothed in pale decumbent simple setae and slightly thicker, white scale-like setae on laterotergites. Antennae with semidecumbent, spine-like setae, AI with several black spines. Legs with black spines. Frons (both sexes) and AI (males) sometimes with long white setae (punctulata). Head mostly smooth, pronotum rugulopunctate, scutellum rugulose, hemelytra finely punctate. Structure: head (Figs 3, 11A– C): Short, transverse; vertex broad, posterior margin not carinate; frons bulbous, extending beyond clypeus; genae tall, approximately twice height of eye;



579

Table 1. Host plant records for the Halticini Genus

Species

Host plant

Family

Reference

arnoldii arnoldii

Helichrysum arenarium Helichrysum arenarium

Asteraceae Asteraceae

Kerzhner, 1964a Putshkov, 1961

dorsalis freyi kirshbaumi longicornis rugicollis

Alyssum calycinum Agropyron sp. Artemisia sp. (Seriphidium) unknown sp. Medicago sp.

Brassicaceae Poaceae Asteraceae Apiaceae Fabaceae

Seidenstücker, 1959 Kerzhner, 1964a Kerzhner, 1964a Kerzhner, 1964a Kerzhner, 1964a

punctulata

Elymus junceus

Poaceae

Kerzhner, 1964a

jakowleffi jakowleffi jakowleffi

unknown sp. unknown sp. Crimea sp.

Poaceae Poaceae Poaceae

Kerzhner, 1964a Wagner, 1973 Wagner, 1973

senecionus

Senecio laxus Leysera tenella Ursinia sp.

Asteraceae Asteraceae Asteraceae

Schwartz et al., 2008 Schwartz et al., 2008 Schwartz et al., 2008

schwartzi

Dampiera incana incana

Goodeniaceae

Tatarnic, 2009

sordidus

Rhanterium epapposum

Asteraceae

Linnavuori, 1986

bleusi cilix lateralis lurensis osellai osellai pygmaeus schmidti seidenstueckeri seidenstueckeri seidenstueckeri tauricus tauricus tauricus tauricus tomasi tomasi tomasi tristis

Stipa tenacissima unknown sp. unknown sp. unknown sp. Lotus sp. Trifolium sp. unknown sp. unknown sp. unknown sp. Teucrium sp. Trifolium sp. Festuca beckeri Festuca sulcata Festuca taurica Poa sp. Festuca sp. Stipa sp. Stipa sp. Anthemis sp.

Poaceae Poaceae Poaceae Poaceae Fabaceae Fabaceae Poaceae Poaceae Asteraceae Lamiaceae Fabaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Asteraceae

Wagner, 1973 Wagner, 1973 Wagner, 1973 Wagner, 1973 Tamanini, 1975 Tamanini, 1975 Wagner, 1973 Franz & Wagner, 1961 Linnavuori, 1984 Linnavuori, 1984 Linnavuori, 1984 Kerzhner, 1964b Kerzhner, 1964b Kerzhner, 1964b Kerzhner, 1964b Tamanini, 1975 Tamanini, 1971 Tamanini, 1976 Linnavuori, 1992

nitidus

unknown sp.

Poaceae

Wagner, 1973

Cassisi

Coopernookia strophiolata

Goodeniaceae

Tatarnic, 2009

schuhi

Goodenia amplexans Scaevola basedowii Scaevola ovalifolia

Goodeniaceae Goodeniaceae Goodeniaceae

Tatarnic, 2009 Tatarnic, 2009 Tatarnic, 2009

apterus apterus apterus beganus beganus insularis insularis intermedius intermedius luteicollis luteicollis luteicollis macrocephalus macrocephalus pusillus pusillus saltator saltator saltator tibialis tibialis tibialis

Vicia striata Vicia sp. Ononis sp. Phaseolus vulgaris ‘rotten oak’ cucumber Ipomoea sp. Anemone canadensis Clematis ligusticifolia unknown sp. unknown sp. Clematis vitalba Carduncellus sp. Clematis sp. Ononis sp. Galium verum cucumber Calendula sp. Althaea sp. Ipomoea sp. Phaseolus sp. unknown sp.

Fabaceae Fabaceae Fabaceae Fabaceae Fagaceae Cucurbitaceae Convolvulaceae Ranunculaceae Ranunculaceae Poaceae Fabaceae Ranunculaceae Asteraceae Ranunculaceae Fabaceae Rubiaceae Cucurbitaceae Asteraceae Malvaceae Convolvulaceae Fabaceae Cucurbitaceae

Hoberlandt, 1956 Wagner, 1973 Wagner, 1973 Alayo, 1974 Henry & Smith, 1979 Carvalho, 1956 Carvalho, 1956 Knight, 1968 Knight, 1968 Ehanno, 1960 Josifov, 1974 Wagner, 1973 Wagner, 1973 Wagner, 1973 Göllner-Scheiding, 1972 Wagner, 1973 Wagner, 1973 Wagner, 1973 Wagner, 1973 Carvalho, 1956 Carvalho, 1956 Carvalho, 1956

hesperius hesperius hesperius hesperius hesperius hesperius hesperius hesperius hesperius hesperius hirtus hirtus

Agropyron cristatum Rosa arkansana unknown sp. Koeleria cristata Poa secunda Stipa comata Stipa williamsi Hordeum sp. Triticum sativum wheat (Triticum sp.) wheat (Triticum sp.) wheat (Triticum sp.)

Poaceae Rosaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae Poaceae

Kelton, 1980 Kelton, 1980 Fuxa & Kamm, 1976 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939 Mills, 1939

Anapomella Anapus

Barbarosia Chorosomella

Compositocoris

Dampierella Dasyscytus Dimorphocoris

Euryopicoris Goodeniaphila Goodeniaphila

Halticus

Labops


580


Table 1. Continued Genus

Species

Host plant

Family

Reference

bractatus bractatus bractatus bractatus

Bryonia sp. Cucurbita moschata Ipomoea batatas Fraxinus sp.

Cucurbitaceae Cucurbitaceae Convolvulaceae Oleaceae

Linnavuori, Maldonado, Maldonado, Linnavuori,

alboornatus alboornatus frontosus macrotrichus macrotrichus oregonensis oregonensis trispiculus

‘native spp.’ Agropyron sp. Psathyrostachys juncea Festuca sp. Carex sp. Agropyron spiculatum Festuca idahoensis Aremisia terrae-albae

Poaceae Poaceae Poaceae Poaceae Cyperaceae Poaceae Poaceae Asteraceae

Bykov, 1971 Kerzhner & Yachevskii, 1967 Drapolyuk & Kerzhner, 2000 Bykov, 1971 Bykov, 1971 Schuh & Lattin, 1980 Schuh & Lattin, 1980 Drapolyuk & Kerzhner, 2000

bivittatus brevis brevis brevis coriaceus coriaceus coriaceus coriaceus coriaceus coriaceus funestrus funestrus medvedvi melas melas minimus proserpinae proserpinae proserpinae saltator saltator saltator saltator scorzonerae scorzonerae vittipennis vittipennis vittipennis vittipennis vittipennis

Artemisia sp. Campanula rapunculoides Campanula rapunculoides Campanula sp. Quercus pedunculata Quercus sp. not specified Tanacetum sp. Achillea sp. Centaurea sp. Artemisia gigantea Artemesia stelleriana Serratula sp. Centaurea sp. Centaurea sp. near Cousinia or Scorzonera Carduus spp. Scolymus spp. Silybum spp. not specified not specified not specified not specified Scorzonera tau-saghyz Takhtajaniantha pusilla Achillea sp. Anthemidae sp. Chrysanthemum sp. Ononis spinosa Chrysanthemum leucanthemum

Asteraceae Campanulaceae Campanulaceae Campanulaceae Fagaceae Fagaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Poaceae Asteraceae Poaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Fabaceae Asteraceae

Kerzhner, 1964a Göllner-Scheiding, 1972 Wagner, 1973 Hoberlandt, 1963 Ehanno, 1965 Ehanno, 1960 Göllner-Scheiding, 1972 Wagner, 1973 Wagner, 1973 Wagner, 1973 Kerzhner, 1978 Kerzhner, 1978 Kerzhner, 1964a Seidenstücker, 1962 Wagner, 1973 Drapolyuk & Kerzhner, 2000 Wagner, 1973 Wagner, 1973 Wagner, 1973 Ehanno, 1960 Göllner-Scheiding, 1972 Gravestein, 1978 Hoberlandt, 1956 Drapolyuk & Kerzhner, 2000 Drapolyuk & Kerzhner, 2000 Göllner-Scheiding, 1972 Hoberlandt, 1956 Göllner-Scheiding, 1972 Göllner-Scheiding, 1972 Wagner, 1973

doriae parallela parallela passerinii

Plantago albicans Potentilla sp. Potentilla sylvestris Thapsia garganica

Plantaginaceae Rosaceae Rosaceae Apiaceae

Wagner, 1973 Wagner, 1973 Kerzhner, 1964A Wagner, 1973

maculatus maculatus ruffoi ruffoi ruffoi zorzii

Teucrium chamaedrys Teucrium chamaedrys Astragalus siculus nebrodensis Prangos sp. Astragalus siculus nebrodensis Globularia sp.

Lamiaceae Lamiaceae Fabaceae Apiaceae Fabaceae Globulariaceae

Ribes, 1984 Wagner, 1973 Tamanini, 1960 Tamanini, 1960 Wagner, 1973 Wagner, 1973

flavomarginatus

not specified

Juncaceae

Wagner, 1973

atrocoeruleus niger niger cicadifrons erythroleptus leucocephalus luridus luridus luridus

Peucedanum oficinale Peucedanum oreoselinum Peucedanum palustre Cistus sp. Calamintha nepeta Campanula spp. Calicotome villosa Jasione montana Jasione montana

Apiaceae Apiaceae Apiaceae Cistaceae Lamiaceae Campanulaceae Fabaceae Campanulaceae Campanulaceae

Wagner, 1973 Wagner, 1973 Wagner, 1973 Wagner, 1973 Wagner, 1973 Göllner-Scheiding, 1972 Pericart, 1965 Göllner-Scheiding, 1970 Göllner-Scheiding, 1972

luridus oberthuri obscurus obscurus raimondoi luridus

Jasione montana Jasione montana Calicotome villosa Jasione montana Elaeoselinum asclepium Jasione montana

Campanulaceae Campanulaceae Fabaceae Campanulaceae Asteraceae Campanulaceae

Wagner, 1973 Wagner, 1973 Pericart, 1965 Wagner, 1973 Carapezza, 1991 Wagner, 1973

Microtechnites 1984 1969 1969 1984

Myrmecophyes

Orthocephalus

Pachytomella

Plagiotylus

Schoenocoris Strongylocoris

Strongylocoris

eyes rounded and substylate, caudally directed, laterally not touching anterolateral margins of pronotum. Antennae (Figs 3, 11A–C): antennal insertion in line with ventral margin of eye; thin, slightly longer

than body in males, in females shorter than body; AI short, swollen in punctulata. Thorax (Figs 3, 11A, C–E): pronotum rectangular, slightly broader posteriorly, pronotal collar indistinct, callosite region weakly



581

Figure 11. Scanning electron micrograph images of Barbarosia punctulata (male). A, head and thorax, dorsal view; B, head, anterior view; C, head and thorax, lateral view; D, external efferent system of the metathoracic scent gland (MTG), detail; E, MTG efferent system; F, tarsus.

defined, lateral margins deflexed, humeral angles rounded, posterior margin weakly concave medially; mesoscutum not visible; scutellum small and triangular; metathoracic spiracle large, exposed, with evaporative bodies posteriorly and dorsally; MTG efferent system broad, swollen, triangular, evaporative area extending to metathoracic spiracle, ostiole prominent, directed laterally, peritreme irregularly rounded. Hemelytra (Figs 3, 11A): coleopteroid, shelllike; clavus and corium fused without subdivided appearance; posterior margins convex; elongate with abdominal tergites VII-IX exposed. Legs (Figs 3, 11F): metafemora incrassate; hindlegs elongate. Male genitalia (Fig. 12A–C): pygophore conical; both parameres of roughly equal size; left paramere L-shaped, weakly concave, apex of apophysis hooked, sensory lobe swollen; right paramere clubbed,

concave, club directed laterally; phallotheca elongateoval; ductus seminis relatively long and slender, with flexible ribbing ending prior to ductus seminis; secondary gonopore sclerotized, horseshoe-shaped basally, opens into less sclerotized, textured hood; endosoma with single, large, elongate S-shaped spicule, basally twisted and bifurcate. Female genitalia (Fig. 12D, E): sclerotized rings of DLP anteriorly narrowed and subcontiguous medially, highly convoluted and heavily sclerotized, narrow, posteriorly recurved forming sinuate short U-shaped canal, posterior margin of VLP sclerotized; posterior wall of bursa copulatrix simple, weakly sclerotized laterally, without processes, medially membranous; vestibular region slightly tumescent, bilaterally symmetrical, opening bordered by short, narrow paired sclerites.


582


Figure 12. Male and female genitalia of Barbarosia punctulata. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, first gonapophyses, ventral view; F, posterior wall.

Diversity and distribution: Barbarosia includes two species and is restricted to eastern Turkey and the Ural Mountains in Russia. Included species: Barbarosia declavata (Seidenstücker, 1962) Turkey Barbarosia punctulata (Reuter, 1901)* Russia: Ural Mts.; Turkey Biology and host plant associations: Barbarosia punctulata has been collected from a single species of grass (Elymus junceus) (Kerzhner & Yachevskii, 1967; Wagner, 1973; Schuh, 1995) (Table 1). Remarks: The biology of Barbarosia is poorly known. It is most similar to Euryopicoris, sharing punctate, coleopteroid, shell-like forewings. Additionally, only in these two genera does the evaporative area of the MTG extend upward nearly to the dorsum (24-2). The apparent proclivity for grass-feeding by Barbarosia and Euryopicoris is also suggestive of a close relationship. However, the male aedeagus and female DLP are unique in Barbarosia, and these characters alone are deserving of genus-group status.

CHOROSOMELLA HORVÁTH (FIGS 3, 13–14) Chorosomella Horváth, 1906: 545 (gen. nov.; type species: Chorosomella jakowleffi Horváth, 1906 by monotypy); Carvalho, 1959: 283 (cat.); Kerzhner,

1962: 385 (tribal placement); Wagner, 1973: 107 (description); Schuh, 1995: 46 (cat.) Diagnosis: Chorosomella is distinguished from all other halticines by the following combination of characters: gracile body; elongate appendages; rectangulate head; frons with conical projection; vertex with midline sulcus; bucculae narrow; eyes distant from anterior margin of pronotum; AI greatly elongate and thick; labium short, reaching mesocoxae; MTG external efferent system obsolete; pretarsi with pulvilli; single elongate endosomal spicule. Redescription: Male macropterous, female micropterous. Coloration (Fig. 3): pale yellow-brown. Surface and vestiture (Figs 3, 13A–D): smooth, almost glabrous. Antennae clothed in very short adpressed spinose setae, becoming most dense on AII through AIV; femora with very short, adpressed spinose setae primarily on dorsal surface; tibiae with slightly longer, semi-erect spine-like setae, shorter than leg width. Structure: head (Figs 3, 13A, B): rectangulate, elongate, length greater than head height; vertex flat with thin medial longitudinal sulcus, posterior margin not carinate; frons with conical projection, extending beyond clypeus; posterior margin of eyes removed from anterior margin of pronotum; genae height subequal to eye height; mandibular and maxillary plates small; bucculae extremely narrow; antennae (Figs 3, 13A, B): inserted at eye midheight; elongate, slender, longer



583

Figure 13. Scanning electron micrograph images of Chorosomella jakwoleffi (male). A, body, lateral view; B, head and thorax, lateral view; C, meso- and metathorax, lateral view; D, meso- and metathorax, detail; E, tarsus.

than body; AI thickest, longer than head and pronotum combined; AII approximately twice as long as AI; AIII slightly longer than first; AIV short. Labium (Fig. 13A, B): short, nearly reaching mesocoxae. Thorax (Figs 3, 13A–D): pronotum narrow, trapezoidal, posteriorly only slightly wider than anterior margin, nearly flat, collar absent, callosite region distinct, lateral margins rounded, posterior margin medially concave; mesoscutum elongate; scutellum short, apex blunt, rounded; metathoracic spiracle a thin slit, with evaporative bodies along posterior margin, MTG external efferent system obsolete. Hemelytra (Fig. 3): males macropterous, costal margin concave, becoming very narrow towards apex; membrane with single cell, apex reaches abdominal tergite VII, minor cell obsolete; females micropterous, hemelytra translucent; without cuneal fracture or membrane; only reaching abdominal tergite III. Legs (Figs 3, 13A, B, E): very long,

cylindrical, thin; pretarsi with fleshy pulvilli. Abdomen (Fig. 1): elongate in both sexes. Male genitalia (Fig. 14A–C): pygophore elongate, posterior margin laterally with pair of caudally projecting processes; parameres roughly equal in size; left paramere narrow, weakly bent, weakly concave, apex of apophysis bifid; right paramere weakly concave, without apical apophysis; phallotheca elongate, narrow and simple, apically slender; ductus seminis elongate, very thin; secondary gonopore weakly sclerotized, opening into elongate tubular spine-like spicule, reaching apex of phallotheca. Female genitalia (Fig. 14D): sclerotized rings thin, elongate, ribbon-like, very weakly sclerotized. DVP and VLP membranous, without sclerotization; posterior wall of bursa copulatrix entirely membranous, without processes or modifications vestibular region symmetrical. Vestibular region symmetrical, very weakly swollen, with


584


Figure 14. Male and female genitalia of Chorosomella jakowleffi. A, aedeagus; B, left paramere; C, right paramere; D, vestibulum; E, bursa copulatrix, dorsal view. Abbreviations: SC, seminal conceptacle; SR, sclerotized ring.

margins wrinkled and membranous, converging anteriorly. Diversity and distribution: Chorosomella is comprised of two species from central districts of the Palaearctic region, including the Russian Crimea and Kazakhstan. Included species: Chorosomella horvathi Kiritshenko, 1911 Kazakhstan Chorosomella jakowleffi Horváth, 1906* Russia Biology and host plant associations: Chorosomella jakowleffi is known to breed and feed on grasses (Kerzhner & Yachevskii, 1967; Wagner, 1973; Schuh & Lattin, 1980) (Table 1). Remarks: Chorosomella was previously placed in the Mirinae: Stenodemini (Horváth, 1906) based on superficial similarities, such as the elongate body and appendages, synonymous with many grass-feeding true bugs. Reuter (1910) transferred the genus to the Orthotylini, but Carvalho (1952) subsequently restored it to the Stenodemini. Kerzhner (1962)

first moved Chorosomella to the Halticini, whereas Wagner (1973) regarded the halticines as a subfamily, and erected a new tribe for Chorosomella: the Chorosomellini. Most authors follow Kerzhner (1962), who based his decision on various characters he considered typical of the Halticini: L-shaped left paramere, spoon-shaped right paramere, membranous aedeagus with spicula, and tall genae, and supported by their close relationship with graminoids. In examining Chorosomella we find that some of these character evaluations are disputable; compared to other Halticini the left paramere is much less angular; the secondary gonopore is unlike that of any other Halticini, and the genae are considerably shorter. Additionally, the long, gracile body shape, long thin legs, pale coloration, the unique shape of the head, and the structure of the female genitalia, particularly the sclerotized rings, are also unlike all other Halticini. In our unweighted phylogenetic analysis Chorosomella groups with Dimorphocoris and Compositocoris (Fig. 1) by several homoplasies, including an overall nonblack coloration (1-0), a slightly elongate head (10-1), and the extreme brachyptery of


THE HALTICINI OF THE WORLD females, with the wings reduced to wing pads (37-3). However, under implied weighting this relationship dissolves, with Chorosomella now appearing as sister to Plagiotylus in a clade that also includes Strongylocoris. This clade is supported by the lack of metathoracic evaporative area (24-0) and the extreme reduction of the metathoracic scent gland itself (23-0). Based on the current data it is unclear which of these competing topologies more closely reflects the true placement of Chorosomella.

COMPOSITOCORIS SCHWARTZ, SCHUH & TATARNIC (FIGS 3, 15–16) Compositocoris Schwartz, Schuh & Tatarnic, 2008: 23 (gen. nov.; type species: Compositocoris senecionus Schwartz et al., 2008 by original designation) Diagnosis: Compositocoris is recognized by the following combination of characters: strongly sexually dimorphic, males macropterous and elongate, females ovoid and micropterous; bright green coloration; females with apically stellate bristle-like setae; medial margins of first gonapophyses forming vestibular opening symmetrical and weakly sclerotized. This genus is very similar to Dimorphocoris but is distinguished by the shorter genal height, coloration, and the apically stellate setae of the females. Diversity and distribution: Compositocoris is a monotypic genus, known only from Namaqualand, South Africa. Included species: Compositocoris senecionus Schwartz et al., 2008* South Africa Biology and host plant associations: This species is known from numerous species of asterids belonging to the genera Senecio, Leyesera, and Ursinia (Asteraceae) (Table 1). Remarks: Compositocoris is very similar to Dimorphocoris in many aspects, including overall male and female body form, the presence of paddle-like sclerites apical to the secondary gonopore, and the patches of honeycombed texturing on the head and thorax (2-1: the only unambiguous synapomorphy linking these as sister taxa in our phylogeny). Only the bright coloration, shorter head, and stellate bristle-like setae in the female distinguish Compositocoris. A more thorough examination of Dimorphocoris is necessary to identify the range of morphological variation and to determine if Compositocoris is congeneric; this was

585

beyond the scope of this study, as we only had access to a few species of the speciose Dimorphocoris.

DAMPIERELLA TATARNIC (FIGS 3, 17–18) Dampierella Tatarnic, 2009: 47 (gen. nov.; type species: Dampierella schwartzi Tatarnic, 2009). Diagnosis: Dampierella is superficially similar to Halticus but may be distinguished by the much shorter head height, the presence of a welldeveloped endosoma with a single spine-like spicule, and lack of wing reduction in either sex. The genitalia of both sexes are typical of the Halticini, with the left paramere bearing a thin apical apophysis, the right paramere elongate and clubbed, the female posterior wall lacking inter-ramal lobes, and the vestibulum weakly sclerotized and bilaterally symmetrical. However, unlike in most other halticines, the base of the left paramere is not elongate, a trait shared with another endemic Australian genus, Goodeniaphila. Diversity and distribution: Dampierella is a monotypic genus, with all specimens collected from two localities in north-west Western Australia. Included species: Dampierella schwartzi Tatarnic, 2009* Australia Biology and host plant associations: Dampierella has only been collected from the plant Dampiera incana var. incana R.Br. (Goodeniaceae) (Table 1). Remarks: This is one of two halticine genera endemic to Australia, the other being Goodeniaphila, both of which are found only on the plant family Goodeniaceae. Our phylogeny places these genera as sister taxa, based on several homoplasious characters and one unambiguous synapomorphy (48-2: the base of the right paramere is elongate, whereas that of left paramere is much shorter). A complete description can be found in Tatarnic (2009).

DASYSCYTUS FIEBER (FIGS 3, 19) Dasyscytus Fieber, 1864: 84 (gen. nov.; type species: Dasyscytus sordidus Fieber, 1864 by monotypy); Kirkaldy, 1906: 131 (cat.); Reuter, 1910: 162 (cat.); Oshanin, 1910: (cat.); Carvalho, 1952: 73 (cat.); Carvalho, 1955: 68 (key); Carvalho, 1958: 8 (cat.); Wagner, 1973: 30 (descr.); Schuh, 1995: 47 (world cat.). Kilicanata Seidenstücker, 1956: 66 (gen. nov., type species: Kilicanata pilifera = junior synonym of Dasyscytus: syn. by Wagner, 1960: 99)


586


Figure 15. Scanning electron micrograph images of Compositocoris senecionus (male and female). A, head of female, dorsal view; B, head and prothorax of female, lateral view; C, head of male, dorsal view; D, meso- and metathorax of male, lateral view; E, external efferent system of the metathoracic scent gland, detail; F, dorsum of female, showing apically stellate setae; G, detail of apically stellate seta; H, pygophore. Modified from Schwartz et al. (2008).

Diagnosis: Distinguished from all other Halticini by the presence of setae on the hemelytral membrane in macropterous males. Redescription: Males macropterous, females coleopteroid or macropterous. Macropterous male

3.2–3.5 mm; macropterous female 3.3–3.5 mm; coleopteroid female 2.5–3 mm. Coloration (Fig. 3): mostly dark brown to black, with some orange-brown, yellow-brown, and yellow markings. Macropterous male mostly orange-brown with yellow and tan markings. Surface and vestiture (Fig. 3): macropterous –



587

Figure 16. Male and female genitalia of Compositocoris senicionus. A, detail of secondary gonopore and endosoma; B, aedeagus; C, left paramere; D, right paramere; E, detail of sclerotized ring, dorsal view; F, first gonapophyses, ventral view; G, posterior wall. Modified from Schwartz et al. (2008).

body clothed in long white setae, with shorter brown setae, including on membrane of hemelytron. Coleopterous – with fewer setae, surface glossy. Antennae with white simple setae and several spinelike setae, most prominent on AI. Pronotum weakly rugulose. Structure: head (Fig. 3): transverse, approximately as tall as broad; width slightly wider than anterior margin of pronotum; vertex transversely sulcate, posterior margin carinate, weakly wraps around pronotum with eyes touching pronotum; genae height slightly greater than eye height; mandibular plate somewhat swollen in macropterous male; maxillary plate carinate. Antennae: insertion in line with lower margin of eye; AI weakly swollen, length approximately equal to eye height; AII weakly apically clavate, slightly more than three times AI length. Thorax (Fig. 3): pronotum trapezoidal, anterior weakly declivent in lateral view, collar not visible, callosite region poorly defined, posterior of humeral angles weakly depressed, posterior margin weakly concave; metathoracic spiracle large and exposed, narrowly bordered with evaporative bodies running downwards but not confluent with metathoracic evaporative area; MTG external efferent system tri-

angular, occupies lower third of tergite, ostiole lateral, tongue-shaped, peritreme rounded, directly above ostiole, surrounded by evaporative bodies. Hemelytra (Fig. 3): macropterous – long and parallel-sided, extending beyond apex of abdomen, cuneus long and thin; membrane with two cells. Coleopterous – reduced to shell-like pads without division, posterior margins running diagonally from apex of claval commissure to lateral margin of abdomen, abdominal tergites IV and V partially covered, VI to apex exposed. Legs: long and slender; metafemora not swollen. Abdomen (Fig. 3): elongate and narrow in macropterous males, not surpassing hemelytral margins; rounded in macropterous females, laterally extending beyond lateral hemelytral margins. Coleopterous: rounded pear-shape. Male genitalia (Fig. 19A–C): pygophore conical; left paramere apically folded, sensory lobe short and narrow, apophysis sinuate and tapering, angled downwards, apically bifid; right paramere spoon-shaped, angled laterally; phallotheca apically narrow, apicodorsally slightly compressed; ductus seminis elongate with flexible ribbing; base of secondary gonopore a sclerotized ring; apically irregularly sclerotized, transversely broad-


588


Figure 17. Scanning electron micrograph images of Dampierella schwartzi (male). A, dorsum; B, head, anterior view; C, head and prothorax, lateral view; D, meso- and metathorax, lateral view; E, pygophore, posterior view; F, pygophore, posteroventral view; G, pygophore, lateral view; H, pretarsus. Modified from Tatarnic (2009).

ened; endosoma with weakly sclerotized dentate folds and apical lobe densely covered with fields of short teeth. Female genitalia (Fig. 19D–F): DLP transverse, sclerotized rings indistinct, triangular-oval, lateral margin most strongly sclerotized, lateral margin and

adjacent portion of DLP weakly upturned; posterior wall of bursa copulatrix membranous with thin, weakly sclerotized rods converging dorsally, ventral margin weakly sclerotized; margins of vestibulum symmetrical and very weakly sclerotized.



589

Figure 18. Male and female genitalia of Dampierella schwartzi. A, left paramere; B, right paramere; C, aedeagus; D, bursa copulatrix, dorsal view; E, posterior wall. Modified from Tatarnic (2009).

Diversity and distribution: This monotypic genus is found throughout much of the Mediterranean region, but has not been collected in Syria, Egypt, Italy, and southern France (Wagner, 1973). Included species: Dasyscytus 1864* Mediterranean

sordidus

Fieber,

Biology and host plant associations: Dasyscytus is found in vegetation at the edges of rivers and ditches, with adults collected in April and May (Wagner, 1973). No clear host records exist, although specimens have been collected under Rhanterium epapposum (Asteraceae) in north-east Arabia (Linnavuori, 1986). Remarks: The monotypic genus Dasyscytus differs from all other halticines by the presence of setae on the hemelytral membrane in macropterous individuals (character 39-1), and the distinct folding of the left paramere (Fig. 19B: not coded in this analysis). In our phylogenetic analysis Dasyscytus is sister to Piezocranum, based on the thin hind femora of females (41-0) and the unbroken sclerotization of the apical region of the ductus seminis up to the secondary gonopore (62-1).

DICYPHOPSIS POPPIUS (FIGS 3, 20–21) Dicyphopsis Poppius, 1914: 11 (gen. nov.; type species: Dicyphopsis nigriceps Poppius, 1914 by monotypy); Carvalho, 1952: 72 (cat.); Carvalho, 1955: 58 (key); Carvalho, 1958: 191 (cat.); Cassis, 1986: 164 (discus-

sion of subfamily placement); Kerzhner & Schuh, 1995: 8 (revised subfamily placement, disc.); Schuh, 1995: 47 (world cat.) Diagnosis: Distinguished from all other Halticini by the following combination of characters: small, gracile body; rounded head with large eyes; semitranslucent hemelytra; aedeagus with extremely narrow, convoluted ductus seminis, endosoma with irregular sclerotized mass apical to secondary gonopore, possibly formed by multiple strand-like spicules. Redescription: Small and fragile; macropterous. Coloration (Fig. 3): glossy; head, pronotum, and abdomen dark brown, contrasting with mostly pale antennae, hemelytra, and legs. Hemelytra translucent. Surface and vestiture (Figs 3, 20A–D, G, H): body sparsely clothed in long pale or dark simple setae and regions of very short, dense, adpressed setae. Head mostly smooth with a few long setae, clypeus and labrum more densely covered with shorter setae; posterior of gena with dense short setae. Antenna clothed in short, slender spine-like setae; AI with a few thin spines, slightly longer than width of segment. Pronotum with sparse covering of longer simple setae, collar with short dense pilosity, thoracic pleura with dense, short pilosity, Hemelytron sparsely covered with long decumbent setae. Femora clothed in long, reclining, simple to weakly spinose setae; tibiae with semi-erect spine-like setae. Structure: head (Figs 3, 20A–D): rounded, weakly transverse, posterior margin of vertex weakly convex; wider than tall,


590


Figure 19. Male and female genitalia of Dasyscytus sordidus. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, posterior wall; F, first gonapophyses, ventral view.

wider than anterior of pronotum; gena height approximately equal to eye height; eyes rounded, sessile, not touching anterior of pronotum; frons broadly rounded, bulging. Labium (Fig. 20C, D): extends to metacoxae. Antennae (Figs 3, 20A–D): insertion in front of ventral margin of eye; long and thin, AI slightly longer than eye height. Thorax (Figs 3, 20D, E): pronotum trapezoidal, steep, collar broad and flat, callosite region faintly medially depressed, humeral angles rounded, posteriorly deeply depressed, lateral and posterior margins concave; mesoscutum visible; scutellum thin and tri-

angular; metathoracic spiracle small, with narrow band of evaporative bodies above; MTG efferent system narrow and angled posteriorly, ostiole a vertical slit, peritreme tongue-shaped, running along ventral margin of tergite, bordered above by narrow band of evaporative bodies. Hemelytra (Fig. 3): semitranslucent, lateral margins parallel, cuneus long and narrow, membrane extends beyond abdomen. Legs (Figs 3, 20A, B, F): long and slender; metafemur slightly incrassate; pretarsi without fleshy pulvilli. Abdomen: narrow, parallel-sided, in males slightly broadened at pygophore. Male genitalia (Figs 20G, H,



591

Figure 20. Scanning electron micrograph images of an undescribed species of Dicyphopsis (male). A, head and thorax, dorsal view; B, head, anterior view; C, head, ventral view; D, head and thorax, lateral view; E, external efferent system of the metathoracic scent gland; F, tarsus; G, pygophore, lateral view; H, pygophore, posterior view.

21A–C): pygophore conical; left paramere L-shaped, sensory lobe broad, apophysis short, tapering, with small subapical keel, apically hooked; right paramere longer than left, straight, spoon-shaped; phallotheca elongate-oval, simple; ductus seminis elongate and constricted, with flexible ribbing; secondary gonopore ill defined, weakly sclerotized; endosoma large, with

broad, irregular sclerotized mass near secondary gonopore, possibly made of many long, thin sclerites. Female genitalia (Fig. 21D–F): DLP membranous, sclerotized rings transverse, medially subcontiguous, subrectangulate, very weakly sclerotized; VLP simple, with medial margin sclerotized and dentate; posterior wall of bursa copulatrix weakly sclerotized anteriorly,


592


Figure 21. Male and female genitalia of an unidentified species of Dicyphopsis. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, ventral view; E, posterior wall; F, first gonapophyses, ventral view.

medially with fields of spines, anteriorly with faint bilateral swellings; margins of vestibulum symmetrical with very weak sclerotization. Diversity and distribution: Dicyphopsis includes only one described species from Tanzania (Kilimanjaro), though undescribed specimens matching the original description and considered congeneric have been collected in Zaire and Ghana (Kerzhner & Schuh, 1995). Included species: Dicyphopsis nigriceps Poppius, 1914 Kilimanjaro.

Remarks: The monotypic genus Dicyphopsis was described from a single female specimen of Dic. nigriceps collected in Tanzania (Poppius, 1914). Unfortunately the holotype was destroyed during shipment to G. Cassis (Cassis, 1986; Kerzhner & Schuh, 1995), leaving no existing specimens of this species. Kerzhner & Schuh (1995) identified a number of specimens that they considered congeneric (although not conspecific) collected in Zaire by N. A. Weber in 1948, and Ghana by R. T. Schuh and J. A. Slater in 1967. We have only seen the Ghana specimens, and although the habitus is not typical of the Halticini, we agree with their assertion that the genitalia is of the Hal-


THE HALTICINI OF THE WORLD ticini type, albeit with some minor differences. Most notably, in the aedeagus the ductus seminis is extremely narrow, and the spicule form is unique. In the female however, the margins of the vestibulum are symmetrical, a character consistent throughout the tribe. It is worth noting that what we call Dicyphopsis superficially appears identical to what Linnavuori (1994) identifies as the Orthotylini genus Nycticapsus Poppius (D. Forero, pers. comm.). Both genera were described in Poppius (1914), with neither one illustrated. As we have not seen specimens of Nycticapsus, for the moment we do not know whether Poppius described the same species twice, whether Linnavuori misidentified specimens of Dicyphopsis, or whether these are in fact distinct genera. As the genitalia are clearly of the Halticini rather than the Orthotylini type, we retain Kerzhner & Schuh’s (1995) initial placement of these specimens. Phylogenetic analysis places Dicyphopsis in a clade of primarily African taxa, defined by the presence of paired bilateral pillow-like swellings on the posterior wall (87-1). These taxa also share similar metathoracic scent gland structure, with the peritreme finger-like, angled back and resting along the caudal margin of the metepimeron (25-3), a condition shared only by Piezocranum and Coridromius (could not be determined in Namaquacapsus because of condition of specimen).

DIMORPHOCORIS FIEBER (FIGS 3, 22–23) Dimorphocoris Reuter, 1890: 253 [gen. nov.; type species: Orthlocephalus signatus Fieber, 1861 subsequent designation (junior synonym of Phytocoris gracilis Rambur, 1839)]; Reuter, 1891: 83, 159 (gen. nov., key); Hueber, 1906: 5, 6 (key, descr.); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 784 (cat.); Reuter, 1910: 147 (cat.); Poppius, 1914: 85 (key, descr.); Bergevin, 1916: 61 (dimorphism); Stichel, 1933: 235 (key); Hedicke, 1935: 59 (key); Kiritshenko, 1951: 126 (key); Wagner, 1952: 96, 107 (key, descr.); Carvalho, 1952: 73 (cat.); Wagner, 1954: 221 (key to spp.); Carvalho, 1955: 67 (key); Carvalho, 1958: 8 (cat.); Lindberg, 1956: 56 (disc., key); Wagner, 1961: 50 (diag., key); Kerzhner, 1964b: 964 (diag., key); Wagner & Weber, 1964: 276 (syn., descr., key); Kerzhner, 1964a: 121 (key to spp. from USSR); Wagner, 1965a: 135 (disc., descr.); Wagner, 1965b: 57 (illustrated key); Wagner, 1969: 79 (modified key); Dimorphocoris (Pedetocoris) Kerzhner, 1970: 634 (subgen. nov., diag., disc., key); Wagner, 1973: 57 (descr., key); Linnavuori, 1992: 216 (key to lateralis group); Ehanno, 1994a: 10–37 (illustrated disc. of morphology, incl. MG, FG); Schuh, 1995: 48 (world cat.). Homoeocoris Fuente, 1918: 236 (gen. nov. type species: Orthocephalus tristis Fieber, 1861: syn. by Wagner, 1955: 241); Carvalho, 1958: 8 (cat.)

593

Diagnosis: Recognized by the following combination of characters: female always brachypterous, male macropterous or brachypterous, macropterous individuals elongate and parallel-sided; body brown with yellow-brown and tan markings; frons, pronotum, and thoracic pleura with faint, symmetrical honeycombed texturing; eyes round and bulging, substylate to stylate; posterior margin of vertex with row of coarse black setae; endosoma with sclerites apparently originating from apex of secondary gonopore. Redescription: Dimorphic, males macropterous or brachypterous, females brachypterous. Coloration (Fig. 3): Highly variable, but generally a combination of dark brown to black with yellow, yellow-brown, or tan markings. Surface and vestiture (Figs 3, 22A–E, G, H): body mostly smooth and impunctate, with some bilaterally symmetrical, honeycomb-like texturing on vertex, frons, pronotum and thoracic pleura, these patches sometimes congruent with contrasting colour patterning. Most of body densely covered with short, pilose setae, much less densely covered with longer simple and spine-like setae; sometimes also with long, silvery scale-like setae; posterior of vertex with a transverse row of coarse black setae; antennae and tibiae with short, semi-reclining bristle-like setae, AI typically with several longer, thicker bristles. Structure: head (Figs 3, 22A–D): transverse, wider than tall; gena height equal or greater than eye height; frons mostly flat and sloping, sometimes with shallow medial sulcus continuing onto vertex; vertex sometimes with bilateral depressions adjacent to eyes, posterior margin of vertex straight, not upturned or carinate, weakly concave to weakly convex; eyes round and bulging, substylate to stylate, projecting laterally beyond and not touching anterolateral margins of pronotum, sometimes angled upwards. Antennae (Figs 3, 22B, C): insertion in line with or below ventral margin of eye; long, thin, and cylindrical, always longer in males but generally not longer than body length; AI sometimes swollen, always longer than eye height; AIV shorter than AIII. Thorax (Figs 3, 22A, D, E): pronotum campanulate in macropterous males, rectangular in brachypterous individuals, gently sloping to nearly level, collar thin, often obscured by strongly tumescent callosite region, posterior margin straight to concave; mesoscutum visible in macropterous males, steeply posteriorly declivent; scutellum often medially tumescent in macropterous males, broad and flat in brachypterous individuals; metathoracic spiracle prominent, diagonal, with evaporative bodies; MTG external efferent system tear-shaped, occupying posterior half of metepisternum, ostiole narrow and orientated ventrolaterally, peritreme elongate, weakly depressed,


594


Figure 22. Scanning electron micrograph images of Dimorphocoris gracilis (male). A, head and pronotum, dorsal view; B, head and prothorax, ventral view; C, head, anterior view; D, head and thorax, lateral view; E, thorax, lateral view; F, tarsus; G, pygophore, ventral view; H, pygophore, lateral view.

surrounded by narrow band of evaporative bodies. Hemelytra (Fig. 3): Macropterous male: elongate, parallel-sided, cuneus long and thin; membrane with two cells, extends beyond tip of abdomen. Brachypterous: usually reduced to an undivided pad, sometimes with faint clavus. Legs (Figs 3, 22F): long, particularly in macropterous males; pretarsi without pulvilli.

Abdomen (Fig. 3): elongate in macropterous males, elongate-oval to pear-shaped in brachypterous individuals. Male genitalia (Figs 23A–C, 22G, H): pygophore triangular in lateral view; left paramere L-shaped, sensory lobe rounded and weakly swollen, apophysis tapering, apex often hooked or weakly bifid; apex of right paramere angled and spoon- or



595

Figure 23. Male and female genitalia of Dimorphocoris gracilis. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, first gonapophyses, ventral view; F, posterior wall.

hockey stick-shaped, apex sometimes with a small apophysis; aedeagus fully enclosed within phallotheca; ductus seminis elongate, with flexible ribbing; secondary gonopore sometimes cup-shaped, often weakly sclerotized and ill-defined, sometimes with scale-like texturing; endosoma with either complex, serrate, sclerotized outpocketings (lateralis group) or with single or paired sclerotized, pedunculate processes, often serrate, emanating from dorsal wall of secondary gonopore (Dim. gracilis group). Female genitalia (Fig. 23D, E): sclerotized rings small, widely separated, mostly round to elongate-oval, lateral margins generally not curved upwards but lateral margin of adjacent portion of dorsal labiate plate typically upturned; posterior wall of bursa copulatrix variable, often divided into two bilaterally symmetrical regions of weak sclerotization, one anterior and one posterior, sometimes partly covered with fields of minute spines; vestibular opening symmetrical, with medial margins of first gonapophyses weakly swollen and weakly sclerotized.

Diversity and distribution: With 58 species this is the largest genus of Halticini. Dimorphocoris is found predominantly in the Mediterranean region but is also known to extend into North and East Africa, Central Asia, and Sakhalin Island in the North Pacific (Ehanno, 1991, 1993, 1994a, b; Wagner, 1973; Linnavuori, 1992; Ehanno & Ribes, 1994). The genus is subdivided into two groups, the Dim. gracilis and Dim. lateralis groups.

Included species: Dimorphocoris abutilon Wagner, 1966 France Dimorphocoris albipilis Kerzhner, 1964 Kazakhstan Dimorphocoris alpinus Poppius, 1910 Kenya Dimorphocoris argaeicus Hoberlandt, 1956 Turkey Dimorphocoris asanovae Kerzhner, 1964 Palaearctic region (western) Dimorphocoris atrans Kerzhner, 1970 Russia Dimorphocoris beieri Wagner, 1965 the former Yugoslavia Dimorphocoris bleusei Puton, 1898 Mediterranean Dimorphocoris carayoni Ehanno & Ribes, 1994 France Dimorphocoris cilix Seidenstücker, 1962 Turkey Dimorphocoris concii Tamanini, 1972 Italy Dimorphocoris constantini Ehanno, 1994a France Dimorphocoris debilis (Reuter, 1880) Mediterranean Dimorphocoris distylus Seidenstücker, 1964 Turkey Dimorphocoris dupuisi Ehanno, 1993a France Dimorphocoris durfortae Ehanno & Ribes, 1994 Spain Dimorphocoris eckerleini Wagner, 1965 Mediterranean Dimorphocoris ehannoi Ribes & Ribes’ 2001 Morocco Dimorphocoris fuscus Joakimov, 1909* Palaearctic region (western)


596


Dimorphocoris gallicus Wagner, 1965 France Dimorphocoris goulae Ehanno & Ribes, 1994 Spain Dimorphocoris gracilis (Rambur, 1839)* Algeria; Spain Dimorphocoris josephinae Ehanno & Ribes, 1994 Spain Dimorphocoris lateralis Reuter, 1901 Palaearctic region (western) Dimorphocoris lividipennis Reuter, 1903 Spain Dimorphocoris longiceps Wagner, 1968 Morocco Dimorphocoris lurensis Wagner, 1957 France Dimorphocoris marci Rizzotti Vlach, 1998 Italy Dimorphocoris marginellus (Puton, 1887) Algeria; Morocco Dimorphocoris mariae Linnavuori, 1952 Mediterranean Dimorphocoris matocqui Ehanno, 1993 France Dimorphocoris mongolicus Kerzhner, 1970 Mongolia Dimorphocoris mutatus Seidenstücker, 1964 Italy Dimorphocoris obachi Ehanno & Ribes, 1994 Spain Dimorphocoris osellai Tamanini, 1976 Italy Dimorphocoris pedetes Kerzhner, 1964 Kazakhstan Dimorphocoris pericarti Tamanini, 1972 France; Italy; Spain Dimorphocoris poggi Carapezza, 2002 Italy Dimorphocoris puigmalis Tamanini, 1976 France Dimorphocoris punctiger (Horváth, 1881) Israel; Syria Dimorphocoris putoni (Reuter, 1882) Andorra; France Dimorphocoris pygmaeus Wagner, 1955 France Dimorphocoris remanei Wagner, 1965 Spain Dimorphocoris robustus Wagner, 1957 France Dimorphocoris ruffoi Tamanini, 1971 Italy Dimorphocoris sari Linnavuori, 1992 Greece Dimorphocoris satyriscus (Scott, 1870) Spain Dimorphocoris saulii Wagner, 1965 Italy; Slovenia; the former Yugoslavia Dimorphocoris schmidti (Fieber, 1858) Ukraine Dimorphocoris seidenstueckeri Linnavuori, 1984 Iraq Dimorphocoris servadeii Tamanini, 1982 Italy Dimorphocoris tamaninii Ehanno, 1993 France Dimorphocoris tauricus (Horváth, 1880)* Russia Dimorphocoris tiberghieni Ehanno, 1993 France Dimorphocoris tomasii Tamanini, 1971 Italy Dimorphocoris toros Seidenstücker, 1962 Turkey Dimorphocoris tristis (Fieber, 1861) Spain Dimorphocoris tuatayae Wagner, 1965 Morocco Biology and host plant associations: Most species are thought to feed on grasses (Wagner, 1973) with the majority of host records coming from the family

Poaceae (Table 1). Some species have also been collected from higher level angiosperms, with Dim. osellai collected from unidentified species of Lotus and Trifolium (Fabaceae) (Tamanini, 1976), Dim. tristis found on Anthemis sp. (Asteraceae) (Linnavuori, 1992), and Dim. seidenstueckeri collected from unidentified species of Asteraceae, Teurcrium (Lamiaceae), and Trifolium (Fabaceae) (Linnavuori, 1984) (Table 1). Many species, especially of the Dim. gracilis group, have restricted mountainous distributions, whereas others are more widely distributed in littoral meadows and on the grassy steppes of North Africa (Wagner & Weber, 1964; Ehanno & Ribes, 1994). It is thought the restricted ranges of many species may be the result of the high degree of brachyptery expressed in the genus, particularly in females (Linnavuori, 1992). Remarks: Dimorphocoris is divided into two species groups: the Dim. lateralis group in which both sexes are brachypterous, and the Dim. gracilis group in which males are macropterous whereas females are brachypterous (Linnavuori, 1992). Linnavuori (1992) further subdivided the former into three ‘phylogenetic subgroups’ (although no phylogenetic results were presented). The highly speciose Dim. gracilis group is also comprised of several distinct subgroupings, which are dealt with in detail by Ehanno (1993, 1994a, b) and Ehanno & Ribes (1994). Members of the Dim. gracilis group are most similar in appearance and in genitalic form to the newly described monotypic genus Compositocoris Schwartz et al., 2008: see appendix). In both cases the males are macropterous whereas the females are brachypterous, and the genitalia of both sexes are very similar. In particular, the presence of two paddleshaped sclerotized processes in the endosoma suggests a close relationship. Only the presence of apically stellate bristles on the body of the female and the structure of the posterior wall set Compositocoris apart. Although there is strong evidence to synonymize Compositocoris with Dimorphocoris, we refrain from doing so without examination of more specimens of Dimorphocoris. Although we dissected few specimens of Dimorphocoris, Ehanno & Ribes (1994; Ehanno, 1994a, b) provide detailed illustrations of many species, including both male and female genital structure, which greatly assisted in informing this redescription.

ECTMETOPTERUS REUTER (FIGS 3, 24–25) Ectmetopterus Reuter, 1906: 59 [gen. nov. type species: Ectmetopterus angusticeps Reuter, 1906 by monotypy (junior synonym of Halticus micantulus Horváth, 1905)]; Oshanin, 1910: 802 (cat.); Reuter,



597

Figure 24. Scanning electron micrograph images of Ectmetopterus micantulus (male). A, head and pronotum, dorsal view; B, head and pronotum, anterior view; C, head, ventral view; D, head and thorax, lateral view; E, thoracic pleura, lateral view; F, tarsus; G, pygophore, posterodorsal view; H, pygophore, ventral view.

1910: 147 (cat.); Hsiao, 1942: 253 (key); Carvalho, 1952: 73 (cat.); Carvalho, 1955: 65 (key); Carvalho, 1958: (10 cat.); Schuh, 1995: 52 (world cat.). Diagnosis: Similar to Halticus and Microtechnites, but readily distinguished by its trifurcate left paramere and broad and flat right paramere.

Redescription: Macropterous. Coloration (Fig. 3): generally black or brown, sometimes with brown and orange-brown markings. Surface and vestiture (Figs 3, 24A–H): body smooth and impunctate, with both simple reclining setae and short scale-like setae. Setae on legs and antennae semi-erect and spine-like. Structure: head (Figs 3, 24A–D): transverse, slightly


598


Figure 25. Male and female genitalia of Ectmetopterus micantulus. A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, posterior wall; F, first gonapophyses, ventral view.

broader than anterior of pronotum; vertex with shallow transverse depression, posterior margin thin and carinate; genae height greater than eye height; eye short and tall; frons steeply sloping; clypeus slightly projecting; maxillary plate swollen; buccula thin. Labium (Fig. 24C, D): LI swollen and elongate, approximately as long as genae height; labium reaches metacoxae. Antennae (Figs 3, 24A–D): insertion in line with lower half of eye; longer than body, cylindrical; AI approximately as long as eye height; AII almost 1.5¥ as long as pronotal width; antennae longer than body length. Thorax (Figs 3, 24A, D, E): pronotum trapezoidal, short, steep, collar thin, lateral margins weakly concave, humeral angles rounded, posterior margin thin, weakly declivent, straight to weakly medially cleft; mesoscutum not visible; met-

athoracic spiracle tear-drop shaped, surrounded with thin border of evaporative bodies extending up along posterior margin of mesepimeron; MTG external efferent system tear-drop shaped, angled posterodorsally, ostiole rounded and orientated laterally, peritreme tongue-like, tapered at apex. Hemelytra (Fig. 3): costal margins evenly curved, broader subapically; clavus apically broadened; R + M vein long, extending almost to cuneus; membrane surpasses apex of abdomen. Legs (Figs 3, 24F): metatibiae slightly swollen; pretarsi with pulvilli. Abdomen: parallel-sided, slightly broader in females. Male genitalia (Figs 25A–C, 24G, H): pygophore conical, posterior margin sinuate, deeply concave below left paramere; parameres with long, thick base; left paramere broad, L-shaped and trifurcate, with apex


THE HALTICINI OF THE WORLD of first branching apophysis hooked, sometimes with apophyses slightly twisted; right paramere straight, flat, and broad, extends out of pygophore, apicolateral margins sometimes strongly recurved; phallotheca elongate-oval, tapering to apex; ductus seminis truncate, with flexible ribbing; secondary gonopore basally sclerotized, apically indistinct; endosoma with weakly sclerotized medial structure extending apically from secondary gonopore. Female genitalia (Fig. 25D, E): DLP unmodified; sclerotized rings widely separated, elongate-ovoid, diagonal, tapered laterally and medially, lateral margins slightly upturned; DLP with a thin transverse sclerotized band adjacent to juncture with posterior wall; margins of VLP sclerotized, medially membranous; posterior wall of bursa copulatrix membranous, medially and laterally with linear fields of spines, laterally with paired, swollen tumescences. Vestibulum symmetrical, swollen, and uniformly weakly sclerotized. Diversity and distribution: All six species of Ectmetopterus are found in eastern Asia. Included species: Ectmetopterus bicoloratus Kulik, 1965* comb. nov. East Russia Ectmetopterus comitans Josifov & Kerzhner, 1972* comb. nov. East Russia Ectmetopterus fuscosus Zou, 1985 comb. nov. China Ectmetopterus maculipes Zou, 1985 comb. nov. China Ectmetopterus micantulus (Horváth, 1905)* East Asia Ectmetopterus niger Zou, 1985 comb. nov. China Biology and host plant associations: No information available. Remarks: Based on similarities in male genitalia, herein we transfer to Ectmetopterus five species of Halticus: Ec. bicoloratus, Ec. comitans, Ec. fuscous, Ec. maculipes, and Ec. niger (all comb. nov.). As with Ec. micantulus, all of these species have East Asian distributions and very similar overall structure. The trifurcate left paramere, long and flat right paramere, and structure of the aedeagus clearly place them in this genus. This redescription is based on examination of Ec. micantulus, Ec. comitans, and Ec. bicoloratus. Although we have not seen the three Chinese species of Zou (1985), based on the descriptions and illustrations of the parameres they are congeneric with the above.

EURYOPICORIS REUTER (FIGS 3, 26–27) Euryopicoris Reuter, 1875b: 11 (gen. nov.; type species: Capsus nitidus Meyer-Dür, 1843 (by mono-

599

typy); Stichel, 1933: 235 (key); Hedicke, 1935: 58 (key); Kiritshenko, 1951: 127 (key); Wagner, 1952: 96, 105 (key, descr.); Carvalho, 1955: 66 (key); Euryopicoris Reuter, 1875b: 24 (typogr. error, key); Reuter, 1875b(1): 87; (2): 99 (key, descr.); Reuter, 1891: 60, 158 (descr., key); Hueber, 1906: 4, 11 (key, descr.); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 790 (cat.); Reuter, 1910: 147 (cat.); Carvalho, 1952: 73 (cat.); Carvalho, 1958: 11 (cat.); Wagner & Weber, 1964: 274 (descr.); Wagner, 1973: 51 (descr.); Schuh, 1995: 53 (world cat.). Diagnosis: Distinguished from other genera by the combination of punctate, coleopteroid hemelytra and curved pro- and mesotibiae. Redescription: Both sexes usually coleopterous, males rarely macropterous. 3–4.5 mm. Coloration (Fig. 3): nearly uniform glossy black, sometimes with yellowbrown at apices of femora and antennal segments, hemelytra of macropterous individuals reddishbrown; eyes red. Surface and vestiture (Figs 3, 26A–F, H): surface glossy; head smooth with radiating folds emanating from centre of vertex; pronotum irregularly rugulose anteriorly, punctate posteriorly; scutellum rugulose; hemelytron punctate. Body with sparse distribution of simple setae, mostly on ventral part of head and ventral and lateral surfaces of body; antennae and legs with semi-erect spine-like setae interspersed with longer, thicker spines, particularly on AI and tibiae. Structure: head (Figs 3, 26A–E): transverse, broader than anterior of pronotum; vertex flat, posterior margin thin, weakly bisinuate; eyes small and rounded, substylate; head height approximately three times eye height; frons broadly rounded, confluent with clypeus; buccula thin; labium (Fig. 26D, E): reaches mesocoxae, LI short and thick. Antennae (Figs 3, 26A–E): insertion in front and below ventral margin of eye; short, approximately as long as body; AI swollen, about two times length of eye height; AIII shorter than AIV. Thorax (Figs 3, 26A, B, E–G): pronotum short, trapezoidal and nearly flat, posterior slightly broader than head, collar absent, callosite region weakly tumescent, lateral margins rounded, humeral angles broadly rounded, posterior margin weakly medially cleft; mesoscutum not visible; scutellum transverse; metathoracic spiracle prominent and elongate, surrounded by a thin band of evaporative bodies; MTG external efferent system broad, triangular, and swollen, ostiole vertically orientated, opens laterally, peritreme tear-shaped, orientated vertically, surrounded by evaporative bodies. Hemelytra (Figs 3, 26A): macropterous: extending beyond apex of abdomen; costal margin weakly convex, nearly straight; membrane with single cell. Coleopterous individuals: long, partially covering tergite VI; undi-


600


Figure 26. Scanning electron micrograph images of Euryopicoris nitidus (female). A, dorsum; B, head and pronotum, dorsal view; C, head, anterior view; D, head, ventral view; E, head and thorax, lateral view; F, thorax, lateral view; G, detail of external efferent system of the metathoracic scent gland evaporative bodies; H, tarsus.

vided; lateral margins rounded, posterior margins weakly rounded. Legs (Figs 3, 26H): metafemora slightly incrassate; fore- and mesotibiae distinctly curved and thickened; pretarsi with fleshy pulvilli. Abdomen (Fig. 3): pear-shaped in both sexes, becoming conical at apex. Male genitalia (Fig. 27A–C):

pygophore conical, genital opening narrow, posterior margin weakly sinuate, with shallow concavity below left paramere; both parameres long and thin basally; left paramere with tumescent sensory lobe, apophysis angled upwards, apically hooked; right paramere with angled apical club, apicolateral margin recurved;



601

Figure 27. Male and female genitalia of Euryopicoris nitidus. A, left paramere; B, right paramere; C, aedeagus; D, bursa copulatrix, ventral view; E, opening of vestibulum; F, posterior wall.

phallotheca broad basally, apically tapering; ductus seminis elongate, basally with flexible ribbing, subapically with elongate, weakly sclerotized section lacking flexible ribbing; secondary gonopore rounded, sclerotized, and U-shaped with prised operculum and fine scale-like texturing; endosoma with a single spine-like spicule. Female genitalia (Fig. 27E, F): sclerotized rings widely separated, strongly sclerotized, oval, lateral, and medial margins and adjacent portions of DLP strongly upturned; lateral margins of DLP moderately sclerotized, medially with transverse bilaterally sulcate sclerotized plate (similar to the inter-ramal bridge in Labops), middle of plate a small carinate point projecting into bursa copulatrix; posterior margin of VLP weakly sclerotized; posterior wall of bursa copulatrix simple and plate-like, medially membranous, laterally weakly sclerotized; vestibulum symmetrical, opening bordered by short, narrow paired sclerites.

Diversity and distribution: There are two species of Euryopicoris, with Euryopicoris fennicus known only from Finland and Euryopicoris nitidus broadly distributed across central Europe into Siberia. Included species: Eu. fennicus Wagner, 1954 Finland Eu. nitidus (Meyer-Dür, 1843)* Central and North Europe; Siberia Biology and host plant associations: Euryopicoris nitidus is found in mountain meadows and is said to live on grasses. Adults are found from June to August. This species overwinters as eggs (Wagner, 1973). There is no biological information for Eu. fennicus. Remarks: Euryopicoris is similar to Anapus and Barbarosia in general structure. Although the secondary gonopore of Euryopicoris is most similar to that of Anapus, the lack of prominent sclerotized processes on the female posterior wall, the punctate hemelytra, and


602


Figure 28. Scanning electron micrograph images of Goodeniaphila cassisi (male). A, dorsum; B, head, anterior view; C, lateral view; D, meso- and metathorax, lateral view; E, pygophore, posterior view; F, pygophore, lateral view; G, pygophore, posteroventral view; H, pretarsus. Modified from Tatarnic (2009).

the curved fore- and mesotibiae of Euryopicoris set it apart. Euryopicoris shares punctate coleopteroid hemelytra and a bulging frons with Barbarosia, but the structure of the DLP in Euryopicoris is relatively simple in contrast to the complexity expressed in Barbarosia. Results of both unweighted and implied weighting phylogenetic analyses placed these three genera in a mono-

phyletic clade, which together form the sister group to Myrmecophyes, Labops, and Scirtetellus.

GOODENIAPHILA TATARNIC (FIGS 1, 28–29) Goodeniaphila Tatarnic, 2009: 52 (gen. nov.; type species: Goodeniaphila cassisi Tatarnic, 2009: 53 by original designation).



603

Figure 29. Male and female genitalia of Goodeniaphila cassisi. A, right paramere; B, left paramere; C, aedeagus; D, aedeagus (phallotheca removed); E, bursa copulatrix, dorsal view; F, posterior wall; G, first gonapophyses, ventral view. Modified from Tatarnic (2009).

Diagnosis: Mostly black with dull olive hemelytra; vestibulum weakly sclerotized and mostly symmetrical, except for small sclerotized knob on left margin. Goodeniaphila most closely resembles Strongylocoris, but can be distinguished by the presence of a welldeveloped MTG external efferent system (absent in Strongylocoris). The dull, olive coloration of the hemelytra and the male and female genitalia set it apart from all other Halticini. Diversity and distribution: This genus is known from two species collected in Western Australia, the Northern Territory, and South Australia. Included species: Goodeniaphila cassisi Tatarnic, 2009* Australia

Goodeniaphila schuhi Tatarnic, 2009*

Australia

Biology and host plant associations: Goodeniaphila has been collected exclusively on the plant family Goodeniaceae.

Remarks: Along with Dampierella, Goodeniaphila is endemic to Australia. Both weighted and unweighted phylogenies place these two in a well-supported sister-taxon relationship. The male genitalia of Goodeniaphila is mostly typical of the Halticini – the left paramere bears an apically hooked apophysis, the right paramere is elongate and clubbed, the ductus seminis is ribbed, the secondary gonopore bears scale-


604


like texturing, and the endosoma is well developed and contains several sclerotized spicules distal to the secondary gonopore. However, the base of the left paramere is short (48-2: shared with Dampierella) rather than elongate as in other halticines. The female genitalia is also mostly typical of the tribe – well-developed sclerotized rings with lateral margins and adjacent portion of DLP weakly upturned, posterior wall without inter-ramal lobes – however, the sclerites of first gonapophyses are nearly symmetrical (as in other halticines) but for a minute sclerotized lobe projecting from the left side (Fig. 29G). This lobe is not seen in other halticines but is present in some Orthotylini, albeit in a much more pronounced state (Schaffner & Schwartz, 2008: 41, fig. 19c). For a complete description of Goodeniaphila see Tatarnic (2009).

HALTICUS HAHN (FIGS 3, 30–32) Astemma Latreille, 1829: 199 (nom. nud.); junior homonym of Astemma Lepeletier & Serville, 1825 [Heteroptera: Pyrrhocoridae]; type species: Acanthia pallicornis Fabricius, 1794 = Cicada aptera Linnaeus, 1758 by monotypy; Westwood, 1840: 121 (description); Amyot & Serville, 1843: 284 (description) Halticus Hahn, 1833: 113 (gen. nov.; type species: Acanthia pallicornis Fabricius, 1794 = Cicada aptera Linnaeus, 1758 by monotypy); Burmeister, 1835: 277 (descr.; Spinola, 1850: 43 (key); Fieber, 1858: 312 (key); Fieber, 1861: 69, 281 (key; descr.); Reuter, 1875a: 23 (key); Reuter, 1875b: 86, 89 (key; descr.); Saunders, 1875: 287 (key); Atkinson, 1890: 117 (cat.); Reuter, 1891: 17, 157 (descr.; key); Saunders, 1892: 267 (key; descr.); Distant, 1904: 479 (descr.); Kirkaldy, 1906: 130 (cat.); Hueber, 1906: 2, 42 (key; descr.); Oshanin, 1910: 802 (cat.); Reuter, 1910: 147 (cat.); Poppius, 1914: 84 (key; descr.); Van Duzee, 1916: 211, 374 (cat.); Knight, 1923: 498 (key); Blatchley, 1926: 800 (eastern US species key); Stichel, 1933: 235 (Palaearctic key); Hedicke, 1935 (Palaearctic key); Knight, 1941: 17, 75, 77 (Illinois key; descr.); Hsiao, 1942: 253 (Chinese key); China, 1943: 268 (nomenclature); Zimmerman, 1948: 199 (Hawaiian key); Slater, 1950: 45 (female genitalia); Kiritshenko, 1951: 127 (USSR key); Wagner, 1952: 95, 96 (Palaearctic key; descr.); Carvalho & Leston, 1952: 245 (British key); Carvalho, 1952: 74 (list); Carvalho, 1955: 65 (key); Carvalho, 1956: 51 (key Micronesian spp.); Carvalho, 1958: 12 (cat.); Southwood & Leston, 1959: 244 (British key); Kelton, 1959: 33, Fig. 81 (male genitalia); Wagner, 1961: 48 (diag.; Palaearctic key); Kerzhner, 1964a: 968 (diag.; USSR key); Maldonado, 1969: 57 (diag.; US key); Alayo, 1974: 15 (diag.); Schuh, 1974: 29 (discussion); Wagner, 1973: 6 (descr.; Palaearctic key); Kelton, 1980: 194 (diag.); Henry, 1983: 610

(Western Hemisphere key); Linnavuori, 1994: 5 (African key); Cassis & Gross, 1995: 186 (synonymy; Australian cat.); Schuh, 1995: 53 (synonymy; world cat.). Eurycephala Laporte, 1832: 36 (gen. nov.; Lygaeus luteicollis Panzer, 1805 by subsequent designation Kirkaldy, 1906: 170); Brullé, 1836: 140 (descr.); Spinola, 1840: 191 (descr.); Blanchard, 1840: 140 (descr.); Reuter, 1891: 17 (synonymy); Carvalho, 1958: 12 (cat.) Halticocoris Douglas & Scott, 1865: 478 (unnecessary nom. nov. for Halticus Hahn nec Haltica Illiger [Coleoptera]); Carvalho, 1958: 13 (cat.); Halticiellus Villiers, 1952: 186 (gen. nov.; type species: Halticus tibialis Reuter, 1891 by monotypy); Carvalho, 1958: 12 (synonymy) Diagnosis: Halticus is characterized by a combination of the following: length of first labial segment less than gena height; elongate, flattened right paramere; left paramere with twisted apophysis; scoop-shaped secondary gonopore and vestigial endosoma lacking spicules; large, thin sclerotized rings. Redescription: Usually small and robust, macropterous or coleopteroid. Coloration (Fig. 3): almost all species black, sometimes with paler yellow, orange, red, or brown colouring on head, antennae, and legs, rarely uniformly brown. Surface and vestiture (Figs 3, 30A–E, G, H): variable, generally glossy, sometimes wholly impunctate, in others pronotum is shallowly punctate; frons and vertex with faint creases radiating from centreline. Body generally with two forms of setae – thin and simple and white, flat, and scale-like; antennae and legs with short semi-erect spines and spine-like setae. Structure: head (Figs 3, 30A, B, D): transverse and short, ranging from broader than tall to taller than broad; vertex with transverse sulcus, posterior margin carinate, concave, generally broader than anterior of pronotum; head height variable, usually > two times eye height; eyes sometimes wrapping around pronotum, sometimes touching; eyes tall and short; clypeus narrow, dorsal margin weakly sulcate; mandibular plate depressed; maxillary plate raised and carinate. Labium (Fig. 30D): length variable, LI greatly swollen. Antennae (Figs 3, 30A–D): insertion typically in line with or slightly below lower margin of eye; long, thin, and whip-like; AI slightly thicker than other segments, about as long as eye height. Thorax (Figs 3, 30A, D–F): pronotum trapezoidal (macropterous) to subrectangular (coleopterous), weakly declivent, laterally sloping, collar thin and flat to absent; callosite region ill-defined to obsolete, posterior margin thinly carinate, straight to weakly medially cleft; metathoracic spiracle prominent, nar-



605

Figure 30. Scanning electron micrograph images of Halticus apterus (male). A, head and pronotum, dorsal view; B, head, anterior view; C, antenna; D, head and thorax, lateral view; E, thorax, lateral view; F, detail of external efferent system of the metathoracic scent gland evaporative bodies; G, tarsus; H, pygophore, ventral view.

rowly bordered with evaporative bodies; MTG external efferent system swollen, occupying lower half of metepimeron, ostiole opens laterally, peritreme rounded, orientated posteriorly, surrounded by evaporative bodies. Hemelytra (Fig. 3): macropterous: costal margin weakly curved, broadest above cuneal fracture; clavus longitudinally folded, with lateral

side steeply sloping; corium laterally sloping; membrane and cuneus deflected. Coleopterous: broadly rounded, laterally and posteriorly sloping; claval suture and medial fracture faint, cuneal fracture very short. Legs: long, metafemur greatly swollen; pretarsi with fleshy pulvilli. Male genitalia (Figs 30H, 31A–E): pygophore conical, posterior margin sinuate, with


606


Figure 31. Male and female genitalia of Halticus apterus: A, phallotheca; B, aedeagus, phallotheca removed; C, secondary gonopore, ventral view; D, left paramere; E, right paramere; F, bursa copulatrix, dorsal view; G, posterior wall; H, bursa copulatrix, ventral view.

Figure 32. Female genitalia of Halticus luteicollis. A, bursa copulatrix, dorsal view; B, first gonapophyses, ventral view; C, posterior wall.

broad deep concavity below left paramere; phallotheca simple and elongate-oval; left paramere apically twisted with sinuate apophysis; right paramere longer than left, extends out of pygophore, broad, flat,

sometimes angled laterally at midpoint, apex rounded with a small apicolateral apophysis; ductus seminis attenuate, with flexible ribbing; secondary gonopore partly sclerotized, in form of elongate scoop, nearly


THE HALTICINI OF THE WORLD filling phallotheca; endosoma extremely reduced, without spicules. Female genitalia (Figs 31F–H, 32A– C): dorsal labiate plate mostly membranous; sclerotized rings large, transverse, medially subcontiguous, sclerotized margins thin, lateral margins weakly upturned; VLP weakly sclerotized on posterior margin, laterally sometimes with paired, scimitarshaped sclerotizations, curving laterally and reaching up to dorsal labiate plate; posterior wall of bursa copulatrix variable, for the most part membranous, often with narrow paired sclerotized bands or rods convergent anteriorly; vestibulum symmetrical, with medial margins of first gonapophyses swollen outwards and sclerotized, vestibular opening rounded and strongly sclerotized. Diversity and distribution: Halticus has 16 species and a mainly Holarctic distribution, with most species confined to the Palaearctic. A single species, Halticus tibialis, is found in Africa, South-East Asia, and the south-west Pacific. Included species: Halticus apterus (Linnaeus, 1758)* Palaearctic Halticus asperulus Horváth, 1898 Armenia Halticus beganus Linnavuori, 1984 Iraq Halticus darbandikhaus Linnavuori, 1984 Iraq Halticus henschii Reuter, 1888 Italy Halticus insularis Usinger, 1946 Mariana Is.; Guam; Korea Halticus intermedius Uhler, 1904 USA Halticus luteicollis (Panzer, 1804)* Europe; Mediterranean Halticus macrocephalus Fieber, 1858 Mediterranean Halticus major Wagner, 1951 Europe Halticus obscurior Kerzhner & Muminov, 1974 Asiatic Russia Halticus puncticollis Fieber, 1870 Montenegro Halticus pusillus (Herrich-Schäeffer, 1835)* Palaearctic Halticus rugosus Reuter, 1894 Syria Halticus saltator (Geoffroy, 1785) Europe Halticus tibialis Reuter, 1891* Africa; South-East Asia, South Pacific Biology and host plant associations: Limited host records exist for Halticus. Halticus apterus and H. beganus have both been collected from the family Fabaceae (Vicia striata and Phaseolus vulgaris, respectively). Halticus beganus has also been collected from rotten oak (Fagaceae) (Henry & Smith, 1979). Halticus pusillus is said to live on and beneath Galium verum (Rubiaceae) (Wagner, 1973) (Table 1).

607

Remarks: Most recent definitions of Halticus are broad, inclusive of great morphological heterogeneity (e.g. Wagner, 1973), and pay little regard to genitalic features. This is the first study to account for the morphological diversity exhibited across species of Halticus, and to examine this in a phylogenetic context. As a consequence we have subdivided Halticus sensu lato into three genera. Based primarily on male and female genitalic characters, and supported by our phylogeny, we remove ten species examined that do not follow the model of the type species, H. apterus (left paramere L-shaped with apically curved apophysis; ductus seminis short; secondary gonopore elongate and scoop-shaped, endosoma obsolete and without spicules; sclerotized rings large and subcontiguous). H. bractatus, H. canus, H. chrysolepis, H. minutus, and H. spegazzini are removed, and along with both species of Cafayatina, form the resurrected genus Microtechnites. Similarly, the Asian species H. bicoloratus, H. comitans, H. maculipes, H. niger, and H. fuscous are all transferred to Ectmetopterus. We have not seen all species of Halticus, and it is therefore possible that with further examination other species may yet be removed from the genus.

LABOPS BURMEISTER (FIGS 4, 33–36) Labops Burmeister, 1835: 279 [gen. nov.; type species: Labops diopsis Burmeister, 1835 by monotypy (junior synonym of Capsus sahlbergii Fallén, 1829)]; Fieber, 1858: 316 (key); Fieber, 1860a: 294; 1861 (key, descr.); Flor, 1860: 448 (key); Thompson, 1871: 432 (key); Walker, 1873: 44 (cat.); Reuter, 1875a: 24 (key); Reuter, 1875b(1): 86 (2):97 (key, descr.); Provancher, 1887: 135 (descr.); Atkinson, 1890: 120 (cat.); Reuter, 1891: 80, 160 (descr., key); Hueber, 1906: 5 (key); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 786 (cat.); Reuter, 1910: 148 (cat.); Van Duzee, 1916: 211 (key); Van Duzee, 1917: 373 (cat.); Knight, 1922: 258 (key, n. spp.); Blatchley, 1926: 797 (key, spp. nov.); Stichel, 1933: 235 (key); Knight, 1941: 81; Slater, 1950: 51 (genitalia); Kiritshenko, 1951: 126 (key); Carvalho, 1952: 74 (cat.); Slater, 1954: 57 (note); Carvalho, 1955: 66 (key); Carvalho, 1958: 18 (cat.); Kerzhner, 1964a: 964 (diag., key); Kelton, 1980: 189 (diag., key); Kerzhner, 1988: 43 (illustrated key to east Asian spp.); Schuh, 1995: 58 (world cat.). Ophthalmocoris Zetterstedt, 1838: 280 gen. nov. (syn. by Herrich-Schäeffer, 1850:166). Diagnosis: Readily distinguished from other Halticini by the following combination of characters: tall head and stylate eyes; hemelytra of brachypterous individuals nearly covering abdomen; aedeagus with mass of thin, needle-like spicules suspended in the membrane anterior to secondary gonopore; posterior


608


Figure 33. Scanning electron micrograph images of Labops sahlbergii (male). A, head and pronotum, dorsal view; B, head, anterior view; C, head and thorax, lateral view; D, meso- and metathorax, lateral view; E, tarsus; F, pygophore, posterior view; G, pygophore, ventral view; H, pygophore, lateral view.

wall of bursa copulatrix with paired inter-ramal sclerites similar to those found in other Orthotylinae; DLP often with heavily sclerotized inter-ramal bridge. Redescription: Coloration (Fig. 4): mostly dark, dull brown to black, usually with yellow, orange, or reddish markings. Head: black with yellow markings, particu-

larly on vertex, along midline, and on genae, mandibular and maxillary plates. Hemelytra black, sometimes with white or yellow along costal margins. Surface and vestiture (Figs 4, 33A–H): clothed in simple setae and pale, scale-like setae. Head smooth and setaceous. Antennae and legs with erect and semi-erect spines, particularly on AI and tibiae. Posterior of pronotum



609

Figure 34. Male and female genitalia of Labops sahlbergii. A, aedeagus, phallotheca removed; B, left paramere; C, right paramere; D, apex of phallotheca, ventral view; E, apex of phallotheca, left side; F, first gonapophyses, ventral view; G, bursa copulatrix; H, posterior wall. Abbreviations: IRB, inter-ramal bridge; IRL, inter-ramal lobe; MP, medial process of ventral labiate plate and ramus.


610


Figure 35. Female genitalia of Labops hesperius. A, bursa copulatrix; B, posterior wall. Abbreviations: IRL, inter-ramal lobe; VLP, ventral labiate plate.

and scutellum rugulose. Hemelytra clothed in simple setae or both simple and scale-like setae. Structure: body elongate-oval. Head (Figs 4, 33A–C): transverse and tall; genae height > two times eye height; eyes stylate, projecting laterally and slightly upward, extending well beyond the anterolateral margins of the pronotum; frons steep, with faint folds radiating from midline; mandibular plate sometimes tumescent, especially males. Labium (Fig. 33C): variable, sometimes reaching metacoxae; LI swollen. Antennae (Figs 4, 33A–C): insertion below eye; shorter than insect; AI swollen, shorter to longer than pronotum, approximately twice as long as eye height. Thorax (Figs 4, 33A, C, D): pronotum campanulate, collar thin and flat, callosite region well defined, posterior margin weakly concave; mesoscutum broad; scutellum small, triangular; metathoracic spiracle small and exposed, thinly surrounded with evaporative bodies; MTG external efferent system broad, swollen and triangular, encompassing lower half of tergite, ostiole prominent and elongate, peritreme tongue-shaped, situated above ostiole in middle of evaporative area, angled rearward parallel to posterior margin of tergite. Hemelytra (Fig. 4): both sexes with brachypterous and macropterous morphs, degree of brachyptery varying across species (Slater, 1954); length variable in brachypterous morphs but generally covering much if not all of the abdomen, cuneus sometimes differentiated, membrane either absent or vestigial; in macropterous morphs hemelytra extend beyond abdomen, margins gently

rounded; membrane with two cells. Legs: long, metafemur not greatly swollen; pretarsi without fleshy pulvilli. Abdomen: elongate-oval. Male genitalia (Figs 33F–H, 34A–E, 36A–D): pygophore simple and conical, ventral margin extending slightly caudad, left ventrolateral margin sulcate below left paramere; left paramere either straight or with obtusely angled apophysis, apophysis weakly curved to strongly angled apically, sensory lobe often prominent; right paramere extends caudally out of pygophore, considerably larger than left paramere, with long base, apex with laterally deflected concave club; apex of phallotheca with serrate, plate-like process of variable size projecting from the left of phallothecal opening; ductus seminis elongate, secondary gonopore sclerotized, vaguely bowl-shaped with a prised operculum, with scale-like texturing along opening margin; endosoma with tight bundle of sclerotized needles, sometimes also with other sclerotized processes, including thin strips of sclerotized teeth. Female genitalia (Figs 34D–G, 35A–B, 36E–G): sclerotized rings large, elongateelliptical, diagonal, medially subcontiguous, lateral margins somewhat upturned; margin of VLP adjoining rami sclerotized, sometimes medially with paired teeth, lateral-most region of VLP joined with rami to form paired, medially projecting, sclerotized processes, sometimes covered with fields of spines; DLP sometimes with heavily sclerotized, bilaterally ventrally pointed inter-ramal bridge, sometimes reduced to paired pincer-like lateral sclerites, a simple band of



611

Figure 36. Male and female genitalia of Labops tumidifrons. A, phallotheca; B, aedeagus, phallotheca removed; C, left paramere; D, right paramere; E, bursa copulatrix, dorsal view; F, inter-ramal bridge; G, posterior wall.

tissue or completely membranous; posterior wall of bursa copulatrix with bilaterally symmetrical interramal lobes of varying complexity projecting from posterior wall; margins of first gonapophyses symmetrical, swollen, and sclerotized. Diversity and distribution: Labops is comprised of 12 species and exhibits a Holarctic distribution. Included species: Labops bami Kulik, 1979 Russia Labops brooksi Slater, 1954 Canada Labops burmeisteri Stål, 1858 Holarctic Labops chelifer Slater, 1954 Canada Labops hesperius Uhler, 1872* Canada; USA Labops hirtus Knight, 1922 Canada; USA

Labops nivchorum Kerzhner, 1988 Russia Labops sahlbergii (Fallén, 1829)* Scandinavia; Siberia (Ural) Labops setosus Reuter, 1891 Russia Labops tumidifrons Knight, 1922* Canada Labops utahensis Slater, 1954* USA Labops verae Knight, 1929 USA Biology and host plant associations: Species of Labops are associated with grasses in arid conditions (Slater, 1954). In North America, L. hesperius has been recorded on Agropyron cristatum, various range grasses, Rosa arkansana (Kelton, 1980), Koeleria cristata, Poa secunda, Stipa comata, Stipa williamsi, Hordeum sp., Triticum sativum (Mills, 1939: in Slater,


612


1954), and wheat (Mills, 1941), whereas L. hirtus has been recorded on wheat in Montana (Mills, 1939). In Yakutia, Vinokurov (1979) recorded the genus on cereals and sedges (Carex stenophylla, Critesion jubatum, and Leymus sp.), and suggested that it may decrease productivity of pastures and hay fields (Table 1). Remarks: Labops has previously been treated as its own tribe, Labopini, owing to various characteristics unique to the genus (Knight, 1923). Slater was the first to note the unusual female genitalia of Labops – both the posterior wall (1950, 1954) and the interramal bridge (1954). Slater (1950) questioned whether the sclerotized inter-ramal tumescences on the posterior wall might be homologous to the interramal lobes (i.e. K-structures) found in other Orthotylinae – the absence of which has been considered a synapomorphy for the Halticini (Schuh, 1974). However, with the discovery of inter-ramal tumescences in both Anapus and Scirtetellus, we consider these to be homologous in origin. Despite the presence of inter-ramal lobes in Labops, Scirtetellus, and Anapus, our phylogeny instead places Labops as sister to Myrmecophyes based on four unambiguous characters, including one synapomorphy (56–1: apex of phallotheca with a prominent flange). The inter-ramal bridge (Kullenberg, 1947; Slater, 1950, 1954) is another interesting structure otherwise absent in the Halticini (although Anapus, Barbarosia, and Euryopicoris possess a sclerotized plate on the DLP, which may be homologous).

MICROTECHNITES BERG (FIGS 4, 37–38) Microtechnites Berg, 1883: 73 [gen. nov.; type species Capsus pygmaeus Berg, 1879 (junior homonym of Capsus (Deraeocoris) pygmaeus Berg, 1878) = Halticus spegazzinii Berg, 1883 by monotypy]; Reuter, 1910: 149 (cat.); Carvalho, 1952: 74 (synonymy with Halticus Hahn); Carvalho, 1958: 13 (cat.); Cassis & Gross, 1995: 186 (cat.; synonymy); Schuh, 1995: 53 (cat.; synonymy). Reinstated status. Cafayatina Carvalho & Carpintero, 1986: 611 (gen. nov.; type species: Cafayatina altigena Carvalho & Carpintero, 1986 by original designation); Schuh, 1995: 46 (cat.). syn. nov. Diagnosis: Microtechnites is recognized by the following combination of characters: macropterous and brachypterous morphs; dark ground colour; moderate distribution of white scale-like setae; eyes sessile; parameres subequal in size; left paramere L-shaped; male endosoma with numerous elongate spicules, often serrate; and, female sclerotized rings widely separated, diagonally orientated. Microtechnites is

similar to Halticus in salient features, but is separated from the latter by the elaborate endosomal spicules and the widely separated sclerotized rings. Redescription: Male and female macropterous and brachypterous morphs, sometimes males macropterous only (e.g. M. bractatus), very small to small, 1.5–3 mm. Coloration (Fig. 4): body mostly dark brown with yellow-brown and yellow markings. Surface and vestiture (Figs 4, 37A–H): body sparsely clothed in semi-erect simple setae intermixed with decumbent, scale-like setae; legs with semi-erect spines. Head mostly smooth, posterior of pronotum weakly rugulose. Structure: head (Figs 4, 37A–C): transverse, elongate in lateral view; posterior margin of vertex sublinear, carinate; head slightly broader than anterior margin of pronotum; frons convex; height of genae approximately equal to height of eye; eyes sessile; maxillary plates sometimes carinate; bucculae narrow; antennae (Figs 4, 37A–C): antennal insertion in front of eye; cylindrical, variable length; AI subequal to eye height, weakly swollen; AII-AIV thinner. Labium (Fig. 37C): reaching metacoxae; LI slightly enlarged; thorax (Figs 4, 37A, C–E): pronotum trapezoidal, collar present and thin, callosite region weakly delineated, posterior margin convex, humeral angles rounded, in lateral view pronotum of macropterous individuals gently sloping, nearly vertical in brachypterous individuals; mesoscutum not visible in both macropterous and brachypterous individuals; scutellum flat, apically rounded; metathoracic spiracle small uniformly bounded by evaporative bodies, sometimes extending to mesepimeral knob; MTG external efferent system broad, pear-shaped, small, confined to medial third of metespisternum, peritreme oval, medially positioned. Hemelytra (Fig. 4): macropterous: lateral margins weakly arcuate; embolium widening towards cuneal fracture; clavus broad, claval commissure elongate; median flexion line half length of claval suture; cuneus transverse; membrane with one cell, minor cell obsolete, membrane extending beyond apex of abdomen. Brachypterous: rounded and shell-like, undivided, posterior margins weakly rounded, wholly or partly covering abdominal tergite VI. Legs (Figs 4, 37F): metafemora elongate, incrassate; metatibiae elongate; pretarsi with fleshy pulvilli. Abdomen: short, lateral margins parallel, slightly broader in female. Male genitalia (Figs 37G, H, 38A–C): pygophore short, conical, genital aperture large, posterior margin symmetrical, concave; left paramere L-shaped, sensory lobe small, apophysis broad, apically arcuate; right paramere subequal to left paramere length, spoon shaped; phallotheca simple; ductus seminis short; secondary gonopore



613

Figure 37. Scanning electron micrograph images of Microtechnites altigena (male). A, head and pronotum, dorsal view; B, head, anterior view; C, head and thorax, lateral view; D, meso- and metathorax, lateral view; E, external efferent system of the metathoracic scent gland; F, tarsus; G, pygophore, posterior view; H, pygophore, ventral view.

with irregular sclerotization; endosoma with several spicules, most spicules elongate, sometimes serrate. Female genitalia (Fig. 38D–F): sclerotized rings moderately sclerotized, widely separated, lateral margins with adjacent portion of DLP weakly upturned; margin of VLP adjacent to rami narrowly

sclerotized, entire, without medial flange; posterior wall of bursa copulatrix subtriangular, weakly sclerotized, anteriorly with paired short digitiform processes; vestibulum symmetrical and weakly sclerotized, anteriorly with paired sclerites converging posteriorly, forming a Y-shape.


614


Figure 38. Male and female genitalia of Microtechnites altigena and Microtechnites canus. Microtechnites altigena: A, aedeagus; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, posterior wall; F, first gonapophyses, ventral view. Microtechnites canus: G, posterior wall.

Diversity and distribution: Microtechnites is restricted to the Western Hemisphere, with most species found in Central and South America. It is the only halticine genus found in the Neotropical region, including its distribution in the Caribbean and the Galapagos Islands (Henry & Wilson, 2004). Only Mi. bractatus is known in the Nearctic region, being widespread in eastern Canada and the USA and extending as far south as Argentina. Microtechnites chrysolepis is endemic to Hawaii and its distributional isolation is significant. We do not regard this disjunction as necessarily anomalous, as the derivation of Hawaii’s fauna from the Western Hemisphere has been postulated for the orthotyline genus Sarona (Asquith, 1995).

Included species: Microtechnites altigena (Carvalho & Carpintero, 1986)* comb. nov. Argentina Microtechnites bractatus (Say, 1832)* Neotropical and Nearctic regions = Rhinacloa citri Ashmead, 1887: 155 (syn. Reuter, 1909) = Halticus minutus Uhler, 1890 nec Reuter, 1884 (junior primary homonym of Halticus minutus Reuter, 1884) = Halticus uhleri Giard, 1892:81 (new name for Halticus minutus Uhler – syn. Parshley, 1915) Microtechnites canus Distant, 1893* (restored status, Henry, 1983: 607) Guatemala; Mexico; Nicaragua Microtechnites chrysolepis (Kirkaldy, 1904)* Hawaii


THE HALTICINI OF THE WORLD Microtechnites inesalti (Carvalho & Carpintero, 1992)* comb. nov. Argentina Microtechnites spegazzinii Berg, 1883* Neotropical region Biology and host plant associations: Host plants have been recorded only for the ubiquitous Mi. bractatus; Maldonado (1969) recorded two cucurbit species as host plants, whereas Linnavuori (1984) recorded Mi. bractatus on sweet potato (Ipomoea batatas) (Table 1). This species, which is commonly known as the garden fleahopper, has been recorded as a pest from a broad range of crops (Henry, 1983; Carpinera, 2001), including solanaceous crops (aubergine, tomato, potato, pepper, tobacco), commercial cucurbits (cucumber, pumpkin, squash) and legumes (alfalfa, beans, clover, cowpeas, peas), as well as beets, cabbage, celery, lettuce, barley, corn, oats, wheat, and cotton. Carpinera (2001) also recorded this species from a great number of weed species, regarded as secondary hosts associated with garden vegetables (Table 1). Microtechnites bractatus is also thought to be zoophytophagous (Carpinera, 2001). It has five instars and overwinters as eggs. Eggs are laid in stems and leaf petioles. Remarks: We have removed Microtechnites from synonymy with Halticus and reinstated it to generic status. In addition, Cafayatina is established as a junior synonym of Microtechnites. Microtechnites is distinguished by its unique male genitalia. In Halticus the aedeagus ends with an elongate scoop-shaped secondary gonopore and the endosoma is vestigial without spicules, whereas in Microtechnites the secondary gonopore opens into a membranous endosoma, which contains several complex spicules and sclerotized serrations. Additionally, the sclerotized rings of Microtechnites are generally smaller and more widely separated, in comparison to Halticus where they are transverse, large, and medially subcontiguous.

MYRMECOPHYES FIEBER (FIGS 4, 39–40) Myrmecophyes Fieber, 1870: 253 [gen. nov.; type species: Myrmecophyes oschannini Fieber, 1870 by monotypy (junior synonym of Diplacus alboornatus Stål, 1858)]; Reuter, 1891: 106 (descr.); Kirkaldy, 1906: 132 (cat.); Oshanin, 1910: 781 (cat.); Reuter, 1910: 148 (cat.); Horváth, 1926: 187 (monograph); Horváth, 1927: 196 (subgen. nov.); Stichel, 1933: 234 (key); Kiritshenko, 1951: 126 (key); Carvalho, 1952: 82 (cat.); Wagner, 1955: 240 (transferred to Halticini); Carvalho, 1955: 79 (key); Carvalho, 1958: 139 (cat.); Bykov, 1971: 871 (diag., key); Wagner, 1973: 4 (descr.); Schuh, 1995: 59 (world cat.).

615

Diplacus Stål, 1858: 183 [gen. nov. (n. preocc. by the nomen nudum Diplacus Rafinesque, 1815 (Ascidiae)]; – (syn. by Reuter, 1890: 252); Reuter, 1875a: 87 (key); 2:100 (descr.); Reuter, 1875b: 25 (key); Atkinson, 1890: 117 (cat.); Carvalho, 1958: 139 (cat.) Diagnosis: Easily recognized by the following characters: strongly myrmecomorphic, with extreme staphylinoidy in females and most males; tall genae in both sexes; hemelytra in macropterous individuals without cuneus, membrane without cells. Redescription: Coloration (Fig. 4): mostly black, sometimes with yellow, red, or orange in parts, sometimes with cream or yellow along the posterior margin of staphylinoid hemelytra, or with mostly pale macropterous hemelytra. Surface and vestiture (Figs 4, 39A–H): surface mostly glossy. Body clothed in simple setae and white, decumbent scale-like setae. Head mostly smooth with a few simple setae; frons with faint rugulose whorls radiating from medial sulcus. Antennae with reclining simple setae, AI with several erect black spines, usually more in males. Pronotum mostly smooth with a few erect simple setae, usually basally and apically rugulose; thoracic pleura clothed in very short, fine setae; scutellum weakly rugulose. Femora with some reclining setae and several black spines dorsally; tibiae with erect spines approximately as long as tibial width, anterior tibiae in males of some species highly spinose. Structure: both sexes strongly ant-mimetic, hemelytra almost always reduced to minute pads, rarely macropterous. Head (Figs 4, 39A–C): transverse, wider than pronotum, taller than wide, genae height > two times eye height; eyes round, substylate, not touching pronotum; posterior margin of vertex weakly concave, not carinate; frons nearly flat, steeply declivent to base of clypeus; clypeus vertical; buccula small and thin. Labium (Fig. 39C): surpasses metacoxae; LI swollen. Antennae (Figs 4, 39A–C): insertion well below ventral margin of eye; long, cylindrical, thin. AI slightly incrassate to significantly enlarged (e.g. My. nasutus), longer than two times eye height. Thorax (Figs 4, 39A, C–E): pronotum quadrate, more campanulate in macropterous males, collar rarely present, callosite region sometimes slightly swollen, lateral margins rounded, posterior margin straight or slightly concave; mesoscutum greatly swollen; scutellum triangular, in staphylinoid morphs anterolateral margins down-curved, apically blunt; metathoracic spiracle prominent, encircled with thin ring of evaporative bodies; MTG external efferent system broad and triangular, evaporative area reaching almost as high as metathoracic spiracle, ostiole elongate, vertically orientated and opening laterally, peritreme broad, tear-shaped, and vertical. Hemelytra (Fig. 4):


616


Figure 39. Scanning electron micrograph images of Myrmecophyes alboornatus (male). A, head and thorax, dorsal view; B, head, anteroventral view; C, head and thorax, lateral view; D, external efferent system of the metathoracic scent gland (MTG) and the metathoracic spiracle; E, detail of MTG evaporative bodies; F, tarsus; G, pygophore, posterior view; H, pygophore, ventral view.

staphylinoid – extremely reduced, only reaching posterior margin of abdominal tergite II, posterior margins rounded, sometimes slightly upturned, sometimes somewhat angular, occasionally claval suture forming a raised carina. Macropterous – clavus present, medial fracture and R + M vein obsolete,

cuneus absent, membrane without closed cells. Legs (Figs 4, 39F): long; protibiae of males in some species expanded (e.g. My. macrotractus); metafemur only somewhat enlarged; pretarsi without fleshy pulvilli. Abdomen (Fig. 4): cylindrical, pear-shaped or globular, usually noticeably rounder in females, basal seg-



617

Figure 40. Male and female genitalia of Myrmecophyes alboornatus. A, aedeagus, phallotheca removed; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, first gonapophyses, ventral view; F, posterior wall.

ments I and II sometimes constricted to form a narrow waist, terminal segments strongly sloping posteroventrally; laterotergites upturned. Male genitalia (Figs 39G, H, 40A–C): pygophore broad, strongly tapered; left paramere weakly bent, sensory lobe often prominent, apophysis straight or curved, apex generally hooked or bifid; right paramere larger than left, sometimes extending caudally from pygophore, apex deflected laterally and spoon-shaped with or without a small apical apophysis; phallotheca with serrate, sclerotized flange (sometimes minute) extend-

ing from left of apex; ductus seminis elongate, with long subapical region more heavily sclerotized and lacking flexible ribbing; secondary gonopore sclerotized and rounded, with partly open operculum, opens ventrally, surface with scale-like texture; endosoma with variable number of sclerotized spicules, generally two or three, apically often with one or two whorl-like sclerites armed with fields of spines. Female genitalia (Fig. 40C–E): sclerotized rings variable in size and shape, ovoid to elongate-elliptical, widely separated, lateral margins and adjacent


618


portion of DLP upturned; DLP and VLP weakly sclerotized and closely adpressed; posterior wall simple and sheet-like, sometimes bilaterally sclerotized; vestibulum symmetrical, with medial margins bordering opening sinuous. Diversity and distribution: Myrmecophyes contains 29 species and has a Holarctic distribution, with a single species (My. oregonensis) restricted to the western USA and the remainder of species found in the Palaearctic region. Included species: Myrmecophyes acutifrons Bykov, 1971 Central Asia Myrmecophyes alacer Horváth, 1926 Transcaucasia Myrmecophyes alboornatus (Stål, 1858)* Palaearctic Myrmecophyes armeniacus Drapolyuk, 1989 Armenia; Turkey Myrmecophyes dubius Bykov, 1971 Central Asia Myrmecophyes ermaki Bykov, 1969 Central Asia Myrmecophyes frontosus Drapolyuk & Kerzhner, 2000 Kazakhstan Myrmecophyes gallicus Wagner, 1976 France; Spain Myrmecophyes geniculatus Reuter, 1894 Central Asia Myrmecophyes heterocerus Horváth, 1926* Armenia Myrmecophyes hirsutiventris Bykov, 1971 Central Asia Myrmecophyes kiritshenkoi Horváth, 1926 Central Asia Myrmecophyes korschinskii Reuter, 1903 Central Asia Myrmecophyes lacteipennis Bykov, 1971 Central Asia Myrmecophyes latus Wagner, 1975 Yugoslavia Myrmecophyes limbatus (Reuter, 1879) Central Asia Myrmecophyes lipskii Bykov, 1971 Central Asia Myrmecophyes macrotrichus Horváth, 1926 Central Asia Myrmecophyes montenegrinus Wagner, 1976 Montenegro Myrmecophyes monticola Horváth, 1926 Central Asia Myrmecophyes muminovi (Kerzhner, 1964) Central Asia Myrmecophyes nasutus Drapolyuk, 1989 Georgia Myrmecophyes nigripes (Reuter, 1879) Central Asia Myrmecophyes nitens Bykov, 1971 Central Asia Myrmecophyes oregonensis Schuh & Lattin, 1980* USA

Myrmecophyes piceus Bykov, 1970 Central Asia Myrmecophyes tibialis Reuter, 1901 Central Asia Myrmecophyes trispiculus Drapolyuk & Kerzhner, 2000 Kazakhstan Myrmecophyes variabilis Drapolyuk, 1989 Azerbaijan Biology and host plant associations: Myrmecophyes is found almost exclusively on grasses. In the USA, My. orgenonensis has been collected from Festuca idahoensis and Agropyron spicatum (Schuh & Lattin, 1980); Kerzhner & Yachevskii (1967) recorded My. alboornatus on Agropyron spp. in Eastern (former) USSR; Bykov (1971) recorded My. macrotrichus on Festuca and Carex spp.; and My. frontosus has been collected from Psathyrostachys juncea (Poaceae) (Drapolyuk & Kerzhner, 2000). A single species has been collected from the family Asteraceae; My. trispiculus is known from the asterid Artemisia terrae-albae (Drapolyuk & Kerzhner, 2000) (Table 1). Remarks: Myrmecophyes is arguably the most convincing ant mimetic genus of Halticini. Based on its mymecomorphy, Carvalho (1952, 1955, 1958) erroneously placed the genus in the Pilophorini. Many characters contradict this assessment (e.g. tall genae, structure of the male and female genitalia) – and in any case several other Halticini are also ant mimics – leading Wagner (1955) and Schuh (1974) to transfer the genus to the Halticini. Schuh & Lattin (1980) provided a brief summary of the genus. The most recent works on Myrmecophyes have been Drapolyuk’s (1989) description of three new species and key to the species of the Caucasus, and Drapolyuk & Kerzhner’s (2000) description of two new species from Kazakhstan. The most comprehensive work remains Bykov (1971), which as Schuh & Lattin (1980) note, omitted nine previously described species from its key and lacks a formal diagnosis. Thus, the most recent diagnosis of Myrmecophyes is that of Wagner (1973), which has been found by us and others (e.g. Schuh & Lattin, 1980) to be both inadequate and inaccurate. Contrary to Wagner, not all species have white markings on the hemelytra, not all species exhibit a macropterous form, the first two abdominal segments are not constricted to form a waist – in fact, only occasionally is the second abdominal segment constricted, whereas in many species the abdomen remains relatively broad at its base – and the opening of the pygophore is not directed downwards. The structure of the ductus seminis and secondary gonopore of Myrmecophyes are most similar to Anapus and Euryopicoris. In particular, in all three genera the distal portion of the ductus seminis lacks flexible sclerotized ribbing. This similarity is reflected



619

Figure 41. Scanning electron micrograph images of Namaquacapsus melanostethoides (male). A, lateral view; B, metatarsus.

Figure 42. Male genitalia of Namaquacapsus melanostethoides. A, aedeagus (phallotheca removed); B, left paramere; C, right paramere.

in the close relationship between these genera (as well as Labops and Scirtetellus) identified in our phylogenetic results. It should be noted that although the two species of Myrmecophyes included in our phylogenetic analysis do not form a monophyletic group, we consider this to be an artefact of our small taxon sample for the genus as well as reflective of the limits of our character dataset.

NAMAQUACAPSUS SCHUH (FIGS 4, 41–42) Namaquacapsus Schuh, 1974: 26 (gen. nov.; type species: Namaquacapsus melanostethoides Schuh, 1974 by monotypy); Schuh, 1995: 62 (cat.)

Diagnosis: Recognized by the following characters: male macropterous (female unknown); robust and elongate; predominantly chestnut-coloured, with corium red at base and apex; impunctate; body clothed in long, erect black setae; tibiae without spines; genae height subequal to eye height; AI slightly swollen, AII apically clavate; pronotum broad; collar flat; pretarsi with minute pulvilli; left paramere L-shaped, with narrow, prominent sensory lobe and long, thin, apically hooked apophysis; right paramere apically spoon-shaped. Redescription: Coloration (Fig. 4): black with rustcoloured hemelytra. Surface and vestiture (Figs 4, 41A, B): head and thorax smooth and glossy, remain-


620


der dull, hemelytra weakly pitted; body clothed with long, black, erect setae; setae on antennae and legs thin and spinose; antennae with short semi-erect setae interspersed with a few long, erect setae; metatibiae without spines. Structure: male macropterous, female unknown. Head (Figs 4, 41): deflexed, short and transverse, broader than anterior of pronotum; posterior margin of vertex raised and carinate; frons broadly convex, steeply declivent; gena height slightly greater than eye height; eyes tall and short, substylate, not touching anterior margins of pronotum; buccula narrow. Labium (Fig. 41A): labium reaches anterior of metacoxa, LI short and swollen. Antennae (Figs 4, 41A): insertion close to and in line with lower margin of eye; thin and cylindrical, shorter than body length; AI slightly swollen, shorter than eye height; AII narrow basally, weakly apically clavate; AIII and AIV thin. Thorax (Fig. 4): pronotum trapezoidal, collar flattened, callosite region obsolete, anterior of humeral angles depressed, posterior margin declivent, weakly rounded, thin and carinate; mesoscutum concealed by pronotum; scutellum strongly convex. Hemelytra (Figs 4, 41A): long; costal margin rounded; clavus steeply declivent laterally; embolium laterally sloping; cuneal fracture deep, cuneus long, laterally rounded; membrane with two cells. Legs (Fig. 41A, B): metafemora only weakly swollen; pretarsi with minute pulvilli. Male genitalia (Fig. 42A–C): parameres elongate, right longer than left; left paramere with narrow, projecting sensory lobe, arm concave, apically bifid; right paramere longer than left, apical club broad, deflected from midline, concave, with apex weakly swollen: phallotheca apically constricted, without lateral projections; ductus seminis long and narrow, with flexible ribbing, subapical portion without ribbing, secondary gonopore sclerotized, curved and dorsoventrally constricted at apex, ventral surface with serrate/scale-like texture; endosoma with weakly sclerotized folds emanating from secondary gonopore, apically with moderately sclerotized serrate and weakly spinose flange. Female unknown. Diversity and distribution: Known only from Namaqualand, South Africa. Included species: Namaquacapsus melanostethoides Schuh, 1974* South Africa Biology and host plant associations: There are no host or ecological data. Remarks: Namaquacapsus is known from only two male specimens. The original description lacked illustrations of the aedeagus, with the endosoma said to be membranous, presumably lacking spicules (Schuh,

1974). Based on our phylogeny, Namaquacapsus belongs to a clade of mostly African taxa that includes Dicyphopsis, Ectmetopterus, Acratheus, and Nanniella, which are united by several characters, including one synapomorphy: the presence of pillow-like swellings on the posterior wall (87-1). As only the male of Nam. melanostethoides is known, the states of this and other female-specific characters are unknown for this species. Additionally, because the legs of the sole specimen we examined obscure our view of the metathoracic scent gland structure, it is uncertain whether or not the peritreme is directed caudally as a thin strip along the posterior margin of the metepimeron (25-2), as is the case in the other clade members.

NANNIELLA REUTER (FIGS 4, 43–44) Nanniella Reuter, 1904: 5 (gen. nov.; type species: Nanniella chalybea Reuter, 1904 by monotypy); Kirkaldy, 1906: 131 (cat.); Reuter, 1910: 148 (cat.); Carvalho, 1952: 76 (syn.); Carvalho, 1958: 61 (cat.); Schuh, 1974: 28 (diag., disc. of tribal placement); Linnavuori, 1975: 48 (disc. of tribal placement, key); Linnavuori, 1994: 9 (descr., key); Schuh, 1995: 62 (world cat.). Diagnosis: Nanniella is distinguished from other Halticini by the following combination of characters: metallic black coloration; uniformly coarsely punctate; left paramere with twisted, somewhat serrate apophysis; right paramere apicolaterally recurved with finger-like apical apophysis. Nanniella is most similar to Acratheus, but can be separated by the presence of deep punctures on the frons and vertex and the shape of the parameres. Redescription: Length: 2.5–4 mm. Colouration (Fig. 4): mostly black with yellow-brown or yellow markings. Surface and vestiture (Figs 4, 43A–G): body with metallic lustre; densely covered with small, slit-like punctures, becoming less dense on abdomen, venter impunctate; frons with deep punctures. Body covered with long, greyish, semidecumbent, simple setae; legs with semi-erect, short spine-like setae. Structure: both sexes macropterous, body elongate and slightly ovate. Head (Figs 4, 43A–D): transverse, slightly broader than tall, broader than anterior of pronotum, height approximately twice eye height; vertex with shallow transverse sulcus, posterior margin slightly upturned and weakly carinate, straight with eyes not touching pronotum; frons almost flat, steeply declivent; clypeus vertical, not projecting forward; buccula small and narrow. Antennae (Figs 4, 43A–B, D): insertion in front and slightly below midpoint of eye, nearly touching eye; long and



621

Figure 43. Scanning electron micrograph images of Nanniella chalybea (male). A, head and thorax, dorsal view; B, head and pronotum, anterior view; C, head and prothorax, ventral view; D, thorax, lateral view; F, external efferent system of the metathoracic scent gland; G, pygophore, ventral view; H, pygophore, lateral view.

thin; AI only slightly thicker than AII, slightly longer than eye height. Thorax (Figs 4, 43A, B, D–F): pronotum trapezoidal, elongate and bulging, collar broad and flat, callosite region poorly defined, lateral margins rounded, posterior of humeral angles weakly depressed, posterior margin wider than head, medially concave; mesoscutum not visible; scutellum flat;

metathoracic spiracle elongate and exposed, without evaporative bodies; MTG efferent system narrow, angled caudally, ostiole nearly ventral, peritreme a thin strip along posteroventral margin of tergite, bordered dorsally by evaporative bodies. Hemelytra (Fig. 4): elongate, lateral margins subparallel, membrane extends beyond abdomen. Legs (Fig. 4): long


622


Figure 44. Male and female genitalia of Nanniella chalybea. A-B, aedeagus; C, left paramere; D-E, right paramere; F, bursa copulatrix, dorsal view; G, first gonapophyses, ventral view; H, posterior wall.

and thin; metafemur not incrassate; pretarsi without fleshy pulvilli. Abdomen (Fig. 43H): parallel-sided, slightly broader at anterior of genital capsule. Male genitalia (Figs 43G, H, 44A–E): pygophore basally broad, short and conical; both parameres long, subequal in length; left paramere elongate, sensory lobe rounded, apical apophysis angled upwards, twisted, dorsally dentate, apically bifid; right paramere projecting slightly out of pygophore, broad, laterally recurved, with curved finger-like apical apophysis; phallotheca simple and elongate; ductus seminis and secondary gonopore broad and dorsoventrally compressed; ductus seminis attenuate, weakly sclerotized, without flexible ribbing; secondary gonopore indistinct, weakly sclerotized, opening into ill-defined

sclerotized channel within endosoma. Female genitalia (Fig. 44F–G): DVP and VLP membranous; sclerotized rings small, thin, and weakly sclerotized, weakly upturned at lateral margins; posterior wall of bursa copulatrix membranous, covered with dense field of minute setae or spines, with very faint lateral swellings; margins of vestibulum weakly swollen, symmetrical, with four small sclerotized patches. Diversity and distribution: All six species of Nanniella share an Afrotropical distribution. Included species: Nanniella alkithoe Linnavuori, 1994 Nigeria


THE HALTICINI OF THE WORLD Nanniella Nanniella Cameroon Nanniella Nanniella Africa Nanniella

chalybea Reuter, 1904* tropical Africa gracilis Linnavuori, 1975 Sudan; pallidiceps Linnavuori, 1973 Zaire palustris Linnavuori, 1975* Equatorial reuteri Poppius, 1914

south-east Africa

Biology and host plant associations: No host information has been collected for this genus. Nanniella alkithoe has been collected at light in savannah forest; Nan. gracilis and Nan. chalybea have both been found in mountain meadows, with the latter also collected in clearings, rain and savannah forests; and Nan. palustris is known from swampy meadows and moist clearings in rain and savannah forests, and is common across western Africa (Linnavuori, 1994). Remarks: Our phylogenetic analysis confirms the sister taxon relationship of Nanniella and Acratheus. As discussed above, both share many similarities, notably the dense, elongate punctures covering the body, the structure of the MTG external efferent system, and the male and female genitalia. This redescription is based on examination of Nan. chalybea and Nan. palustris, as well as review of the literature (Linnavuori, 1994).

ORTHOCEPHALUS FIEBER (FIGS 4, 45–46) Orthocephalus Fieber, 1858: 316 (gen. nov.; type species: Lygaeus brevis Panzer, 1798 by subsequent designation Reuter, 1888: 76); Fieber, 1858: 316 (gen. nov., key); Fieber, 1860b: 291, 1861 (key, descr.); Douglas & Scott, 1865: 429 (descr.); Thompson, 1871: 432 (key); Reuter, 1875b: 24 (key); Reuter, 1875a 1:86; 2:93 (key, descr.); Saunders, 1875: 289 (key); Provancher, 1887: 136 (descr.); Reuter, 1891: 43, 158 (key, descr.); Saunders, 1892: 269 (key, descr.); Hueber, 1906: 3, 15 (key, descr.); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 791 (cat.); Reuter, 1910: 148 (cat.); Van Duzee, 1914: 389 (cat.); Knight, 1923: 498 (key); Blatchley, 1926: 804 (descr.); Stichel, 1933: 235 (key); Hedicke, 1935: 59 (key); Knight, 1941: 75, 81 (key); Hsiao, 1942: 253 (key); China, 1943: 267 (cat.); Slater, 1950: 46 (female genitalia); Kiritshenko, 1951: 127 (key); Wagner, 1952: 96, 103 (key, descr.); Carvalho & Leston, 1952: 245 (key, fig.); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 22 (cat.); Southwood & Leston, 1959: 247 (key); Wagner, 1961: 50 (diag., key); Kerzhner, 1964a: 966 (diag., key); Wagner & Weber, 1964: 267 (descr., key); Wagner, 1973: 41 (descr., key); Schuh, 1995: 63 (world cat.); Namyatova & Konstantinov, 2009 (generic revision, phylogeny).

623

Anapomella Putshkov, 1961: 25 (syn. by Namyatova & Konstantinov, 2009) Oraniella Reuter, 1894: 138 (syn. by Namyatova & Konstantinov, 2009) Diagnosis: Orthocephalus can be recognized by the following combination of characters: head transverse with frons flat to moderately convex; eyes sessile; AI short with two or three mesial spine-like setae, dorsum impunctate, usually covered with robust, bristle-like setae, apical portion of ductus seminis bowl-shaped, strongly dilated and sclerotized, apically funnel-shaped and dorsoventrally compressed, with distinct scale-like sculpturing. Redescription: Coloration (Fig. 4): body dark brown to black, sometimes with yellow to reddish-brown colouring, particularly on legs; hemelytra of brachypterous morphs sometimes pale tan and brown. Surface and vestiture (Figs 4, 45A–H): dorsum impunctate, pronotum, scutellum, and hemelytron sometimes rugulose; body clothed with long, black, bristle-like semi-erect setae; AI with two or three spines; femora sometimes with a few spines, tibiae strongly spinose; some species also densely clothed in white, scale-like setae. Structure: degree of wing polymorphism varies across species, both sexes macropterous or brachypterous. Head (Figs 4, 45A–C): transverse, slightly broader than anterior of pronotum, always narrower than posterior of pronotum in macropterous males; posterior margin of vertex rounded to weakly carinate; frons flat to moderately convex, steeply declivent; gena height slightly greater than eye height; eyes round, substylate. Labium (Fig. 45C): variable, reaching from before procoxae to slightly beyond metacoxae; LI short and swollen. Antennae (Figs 4, 45A–C): insertion close to and in line with lower margin of eye; thin and cylindrical, shorter than body length; AI sometimes somewhat swollen, short, sometimes slightly longer than eye height; AII sometimes apically clavate; AIII and AIV thin. Thorax (Figs 4, 45A–D): pronotum trapezoidal, in macropterous morphs declivent with thin collar, in brachypterous morphs sometimes more rectangular and flat without collar; callosite region present but sometimes ill-defined, lateral margins rounded in brachypterous morphs, posterior margin straight to weakly concave; mesoscutum sometimes concealed by pronotum in brachypterous morphs; scutellum flat to somewhat swollen; metathoracic spiracle conspicuous and exposed, often large, narrowly surrounded with evaporative bodies that extend up to dorsolateral margin of metathorax; MTG efferent system swollen and triangular, extending as high as base of mesocoxae, ostiole vertical, opens laterally, peritreme oval, extending posterodorsally from ostiole. Hemelytra


624


Figure 45. Scanning electron micrograph images of Orthocephalus brevis (male). A, head and thorax, dorsal view; B, head, anterior view; C, head and prothorax, lateral view; D, meso- and metathorax, lateral view; E, tarsus; F, pygophore, dorsal view; G, pygophore, posterior view; H, pygophore, ventral view.

(Fig. 4): macropterous – long and parallel-sided, all divisions present, cuneus long and narrow, membrane with two cells; brachypterous – membrane absent, all remaining divisions faint but present, cuneal fracture very short, posterior margin rounded. Legs (Figs 4, 45E): metafemora moderately swollen; pretarsi without fleshy pulvilli. Male genitalia (Figs 45G, H,

46A–D): pygophore conical, posterior margin weakly asymmetrical, sometimes with thin apical serrate flange (e.g. Orthocephalus modarresi); parameres of roughly equal length; left paramere L-shaped, sometimes with pronounced sensory lobe, apex of apophysis hooked, sometimes bifid; right paramere with spoon-like apex deflected laterally, with small apical



625

Figure 46. Male and female genitalia of Orthocephalus brevis. A, aedeagus; B, secondary gonopore; C, left paramere; D, right paramere; E, posterior wall; F, bursa copulatrix, dorsal view.

apophysis; ductus seminis attenuate, with flexible ribbing, secondary gonopore moderately sclerotized, basally constricted to form dorsoventral bowl shape, apex dorsoventrally compressed and laterally expanded, with scale-like sculpturing primarily on ventral wall, dorsal wall longer than ventral wall; endosoma usually with two or three spinose spicules, rarely with none. Female genitalia (Fig. 46E, F): sclerotized rings widely separated, subtriangular, lateral and medial margins of DLP adjacent to sclerotized rings weakly sclerotized and upturned; VLP divided into anterior band attached to rami and medial plate,

both portions weakly sclerotized; posterior wall of bursa copulatrix a simple, undivided, weakly sclerotized plate; vestibulum symmetrical, margins weakly sclerotized and unmodified.

Diversity and distribution: Orthocephalus includes 23 species and has a Holarctic distribution. One species, Orthocephalus coriaceus, is found both in Europe and eastern North America, whereas all other species are restricted to the Palaearctic region.


626


Included species: Orthocephalus arnoldii (Putshkov, 1961)* Kazakhstan; Ukraine Orthocephalus bivittatus Fieber, 1864 Turkey; Turkestan, East Asia (USSR) Orthocephalus brevis (Panzer, 1798)* Palaearctic Orthocephalus championi Saunders, 1894 Corsica Orthocephalus coriaceus (Fabricius, 1777)* Europe; North America Orthocephalus fulvipes Reuter, 1904* Mediterranean; Asia Minor Orthocephalus funestus Jakovlev, 1881 China; Siberia; north-east Asia; Vladivostok Orthocephalus medvedevi Kiritshenko, 1951 Ukraine Orthocephalus melas Seidenstücker, 1962 Turkey Orthocephalus minimus Drapolyuk & Kerzhner, 2000 Kazakhstan Orthocephalus modarresi Linnavuori, 1997 Iran Orthocephalus proserpinae (Mulsant & Rey, 1852) Mediterranean Orthocephalus putshkovi (Namyatova & Konstantinov) Ukraine; Kazakhstan Orthocephalus rhyparopus Fieber, 1864 south Russia Orthocephalus saltator (Hahn, 1835) Holarctic Orthocephalus scorzonerae Drapolyuk & Kerzhner, 2000 Uzbekistan; Turkmenistan; Kazakhstan Orthocephalus sefrensis Reuter, 1895 Algeria Orthocephalus solidus (Seidenstücker, 1971 Turkey Orthocephalus styx Reuter, 1908 Turkey; Russia; Kazakhstan Orthocephalus tibialis (Reuter, 1894) Algeria; Morocco; Tunisia Orthocephalus tristis (Reuter, 1894) Algeria Orthocephalus turkmenicus Namyatova & Konstantinov, 2009 Turkmenistan; Iran Orthocephalus vittipennis (Herrich-Schäeffer, 1835) Palaearctic Biology and host plant associations: Host records exist for 13 species, all of which have been collected mainly on asterids in the family Asteraceae, with a single species (O. brevis) known only from the Campanulaceae (Hoberlandt, 1963; Göllner-Scheiding, 1972; Wagner, 1973). In addition to feeding on asterids, three species have also been collected off rosids, with O. coriaceus found on oak (Fagaceae) (Ehanno, 1960, 1965), O. vittipenis on Ononis spinosa (Fabaceae) (Göllner-Scheiding, 1972), and O. saltator on Spiraea sp. (Namyatova & Konstantinov, 2009). Orthocephalus saltator has also been collected on unidentified grass (Poaceae) (Ehanno, 1960; Gravestein, 1978), several species of Fabaceae (Reuter, 1891; Saunders, 1892; Tamanini, 1981; Seidenstücker, 1959), as well as Thymus serpyllum (Lamiaceae)

(Reuter, 1891) and Salix sp. (Salicaceae) (Namyatova & Konstantinov, 2009) (Table 1). Remarks: Orthocephalus is superficially similar to Dasyscytus, Pachytomella, and Anapus, but can be readily distinguished by the unique structure of the secondary gonopore. In a recent revision of Orthocephalus, Anapomella Putshkov and Oraniella Reuter were designated junior synonyms of Orthocephalus, based primarily on the secondary gonopore structure (Namyatova & Konstantinov, 2009). This was supported through a phylogenetic analysis, which also proposed a sister relationship between Orthocephalus and Pachytomella. Our phylogeny supports this sister taxon relationship as well, based on two synapomorphies (69-1: opening of secondary gonopore slit-like; 83-1: DLP with pair of sickle-shaped sclerites adjacent to sclerotized rings). Interestingly, although we incorporated the coding of Namyatova & Konstantinov (2009) into our analysis, only the slit-shaped secondary gonopore opening was found to be synapomorphic in both analyses. This appears to be the result of differences in taxon sampling, as Namyatova & Konstantinov (2009) sampled all Orthocephalus, not all of which possess paired sickle-shaped sclerites on the DLP.

PACHYTOMELLA REUTER, 1890 (FIGS 4, 47–48) Pachytoma Costa, 1842: 288 [gen. nov.; type species: Pachytoma minor Costa, 1842, =Phytocoris passerinii Costa, 1842 by monotypy; junior homonym of Pachytoma Svains, 1840 (Mollusca)]. Pachytomella Reuter, 1890: 253 (new name for Pachytoma Costa, 1842); Reuter, 1891: 37, 158 (key, descr.); Hueber, 1906: 3, 27 (key, descr.); Oshanin, 1910: 796 (cat.); Reuter, 1910: 148 (cat.); Stichel, 1933: 235 (key); Hedicke, 1935: 58 (key); China, 1943: 267 (cat.); Wagner, 1952: 96, 101 (key, descr.); Carvalho & Leston, 1952: 245 (key, fig.); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 27 (cat.); Carvalho, 1958: 27 (cat.); Wagner, 1961: 49 (diag., key); Wagner & Weber, 1964: 265 (syn., descr., key); Wagner, 1973: 32 (descr., key); Schuh, 1995: 67 (world cat.). Diagnosis: Recognized by the following combination of characters: head transverse, posterior margin wrapping around pronotum; hemelytra parallel-sided; MTG obsolete. Redescription: Sexually dimorphic, with males macropterous or brachypterous and females brachypterous. Length = 1.5–4 mm. Coloration (Fig. 4): mostly dark brown to black, sometimes with the hemelytra partly to mostly pale, legs usually dark, sometimes



627

Figure 47. Scanning electron micrograph images of Pachytomella passerini (male). A, head and thorax, lateral view; B, head and pronotum, dorsal view; C, head, anterior view; D, meso- and metathorax, lateral view; E, detail of metathorax; F, tarsus; G, pygophore, lateral view; H, pygophore, ventral view.

partly to mostly yellow-brown or yellow. Surface and vestiture (Figs 4, 47A–H): impunctate, hemelytra and posterior of pronotum rugulose. Clothed in long, decumbent, simple setae; setae on antennae and legs spinose and semi-erect interspersed with longer spines; setae on pygophore long and directed caudally. Structure: head (Figs 4, 47A–C): transverse, appear-

ing triangular from above in some species; slightly broader than anterior of pronotum, much broader than tall; genae height greater than eye height; posterior margin of vertex carinate; frons steep; eyes large and round, deflected caudally and touching pronotum; buccula thin; labium (Fig. 47A): LI short and broad. Antennae (Figs 4, 47A–C): insertion adja-


628


Figure 48. Male and female genitalia of Pachytomella passerini. A-B, aedeagus; C, left paramere; D, right paramere; E, first gonapophyses, ventral view; F, sclerotized rings, dorsal view; G, posterior wall.

cent to lower margin of eye; always shorter than body; AI as long as eye height, somewhat thicker than AII. Thorax (Figs 4, 47A, B, D, E): pronotum trapezoidal, becoming subrectangular in brachypterous morphs, short, collar absent, callosite region weakly defined in macropterous morphs, obsolete in brachypterous females, lateral margins rounded, posterior margin approximately equal to head width, straight to concave; mesoscutum visible in macropterous males; scutellum broad in brachypterous morphs; metathoracic spiracle small and narrow; MTG external efferent system obsolete. Hemelytra (Fig. 4): macropterous males – elongate, parallel-sided; claval commissure elongate; cuneus narrow and short; membrane with two cells, extends beyond abdomen. Brachypterous – with or without distinct clavus; without cuneus or membrane; extending partway over abdominal tergite VI. Legs (Figs 4, 47F): short; metafemora incrassate in females; pretarsi without pulvilli. Abdomen (Fig. 4): elongate-oval in males, broader and pearshaped in females. Male genitalia (Figs 47G, H, 48A– D): pygophore broad and conical; posterior margin weakly concave beneath left paramere; parameres long and slender; left paramere narrow, sensory lobe small or absent, apical apophysis sharply curved or angled, apex sometimes broadly hooked; right paramere sometimes projecting from pygophore, apical club elongate, weakly laterally deflected; phallotheca short, basally broad and weakly laterally constricted; ductus seminis attenuate, with flexible

ribbing, subapically sclerotized and lacking ribbing until secondary gonopore; secondary gonopore roughly circular, usually dorsoventrally compressed (less so in Pachytomella passerini) with distinct scalelike texturing; endosoma with or without one or two broad, sclerotized spicules. Female genitalia (Fig. 48E, F): sclerotized rings moderately sized, rectangular, widely separated, lateral margins and adjacent portion of DLP distinctly curved upwards; anterolateral margins of VLP sclerotized; posterior wall undivided and plate-like, mostly membranous with weak sclerotization along anterior margin; opening to vestibulum symmetrical, with lateral margins weakly sclerotized and unmodified. Diversity and distribution: Pachytomella is a European genus of six species, which are almost exclusively found in the Mediterranean region. Included species: Pachytomella alutacea (Puton, 1874) Spain Pachytomella cursitans Reuter, 1905b Spain Pachytomella doriae (Reuter, 1884)* Spain; Tunisia Pachytomella parallela (Meyer-Dür, 1843)* central and north Europe Pachytomella passerinii (Costa, 1842)* Mediterranean Pachytomella phoenicea (Horváth, 1884) eastern Mediterranean



629

Figure 49. Male and female genitalia of Piezocranum simulans. A, aedeagus, phallotheca removed; B, left paramere; C, right paramere; D, bursa copulatrix, dorsal view; E, detail of bursa copulatrix, ventral view; F, posterior wall; G, first gonapophyses, ventral view.

Biology and host plant associations: Limited host and biological records have been published for this genus. Two species for which host records exist are associated with high altitude: Pa. alutacea is found in high mountains under Thymus spp. and other plants, whereas Pa. parallela is found in mountain meadows on Potentilla spp., with adults present from June to August (Wagner, 1973). Pachytomella passerini has been collected on Potentilla sylvestris (Rosaceae) (Kerzhner, 1964a) and Thapsia garganica (Apiaceae) (Wagner, 1973). This species is associated with arid areas, with adults present from May to August. Pachytomella passerini overwinter as eggs (Wagner, 1973). Pachytomella doriae has been collected on Plantago albicans (Plantaginaceae), with adults present in May (Wagner, 1973) (Table 1).

Remarks: Within the Halticini, Pachytomella shares with Chorosomella, Plagiotylus, and Strongylocoris extreme reduction of the MTG, which based on our phylogenetic analysis, has occurred multiple times (character 23-1). As discussed above, Pachytomella is sister taxon to Orthocephalus, a relationship supported both in our analysis and that of Namyatova & Konstantinov (2009). All three genera are easily distinguished by their coloration, body shape, and male and female genitalia.

PIEZOCRANUM HORVÁTH (FIGS 4, 49) Piezocranum Horváth, 1877: 92 (gen. nov.; type species: Piezocranum simulans Horváth, 1877 by


630


monotypy); Atkinson, 1890: 120 (cat.); Reuter, 1891: 33, 157 (descr., key); Hueber, 1906: 3 (key); Kirkaldy, 1906: 130 (cat.); Reuter, 1910: 148 (cat.); Oshanin, 1910: 798 (cat.); Stichel, 1933: 235 (key); Kiritshenko, 1951: 127 (key); Wagner, 1952: 96, 101 (key, descr.); Carvalho, 1952: 74; Carvalho, 1955: 67 (key); Carvalho, 1958: 29 (cat.); Kerzhner, 1964a: 967 (diag., key); Wagner & Weber, 1964: 264 (syn., diag., key); Wagner, 1973: 28 (descr., key); Schuh, 1995: 68 (world cat.). Lamprella Reuter, 1890: 253 (gen. nov.) – (syn. by Horváth, 1889: 327); Lamprella Carvalho, 1958: 29 (cat.). Diagnosis: Recognized by the following combination of characters: macropterous individuals elongate, with transverse depression spanning vertex and frons; endosoma without spicules; DLP divided into weakly sclerotized lateral and medial plates, with paired sclerotized strips adjacent to medial margin of sclerotized rings. Redescription: Sexually dimorphic, with males macropterous and females macropterous or brachypterous. Coloration (Fig. 4): Male reddish-brown, paler on hemelytron; female glossy black. Surface and vestiture (Fig. 4): head smooth and shining, pronotum rugulose, especially along posterior margin, hemelytra of female sometimes glossy with deep punctures. Body of male clothed in golden simple setae; females less setaceous. Structure: head (Fig. 4): transverse; usually wider than tall; macropterous morphs with deep transverse depression spanning across vertex and frons, brachypterous females with at most a shallow depression; posterior margin of vertex raised and carinate; frons broadly rounded in brachypterous females; gena height two to three times eye height; labium: LI swollen, short. Antennae (Fig. 4): insertion below or in line with lower margin of eye; shorter than body, slender; length of AI approximately equal to eye height. Thorax (Fig. 4): pronotum trapezoidal in macropterous morphs, subrectangular in brachypterous females, callosite region obsolete to weakly defined in both sexes, lateral margins rounded, humeral angles rounded, posterior margin concave, more deeply in female; mesoscutum visible from above in macropterous individuals; metathoracic spiracle tear-shaped and exposed, without evaporative bodies; MTG external efferent system swollen, angled posterodorsally, ostiole ventrolateral, peritreme tongue-shaped. Hemelytra (Fig. 4): macropterous – parallel-sided, elongate, extending beyond apex of abdomen; cuneus elongate; membrane with two cells. Brachypterous females – undivided, posteriorly almost straight to subtriangular, sometimes extending partly over abdominal tergite VI. Legs (Fig. 4):

short, somewhat longer in brachypterous morphs; pretarsi without fleshy pulvilli. Abdomen (Fig. 4): narrow in macropterous individuals, broader in females, in brachypterous females pear-shaped. Male genitalia (Fig. 49A–C): pygophore conical; parameres roughly equal in length; left paramere L-shaped, sensory lobe broad, apophysis slender and apically hooked; right paramere slender basally, apical club elongate, weakly deflected laterally, inner margin sinuate; phallotheca unmodified; ductus seminis constricted, with flexible ribbing; secondary gonopore elongate, incompletely sclerotized, dorsoventrally compressed, apically with scale-like texturing; endosoma simple and broad, without spicules. Female genitalia (Fig. 49D–F): sclerotized rings widely separated, ovate, weakly sclerotized; lateral margins of DLP upturned and sclerotized, closely appressed to rings, remainder of DLP and VLP membranous; posterior wall mostly membranous, with faintly sclerotized band along posterior margin; opening to vestibulum symmetrical, membranous, and unmodified. Diversity and distribution: Piezocranum is composed of three species, all of which are found almost exclusively in the Mediterranean region, with the range of Piezocranum simulans also extending into the former Yugoslavia, Bulgaria, and Romania. Included species: Piezocranum corvinum Puton, 1895 Syria; Turkey Piezocranum seminulum Horváth, 1898 Spain Piezocranum simulans Horváth, 1877* southern Europe, Macedonia Biology and host plant associations: No biological or host plant information has been recorded for this genus. Remarks: This redescription is based on a review of the literature, particularly Wagner (1973), and examination of Pi. simulans. Our phylogeny places Piezocranum as sister to Dasyscytus, although this relationship is not well supported.

PLAGIOTYLUS SCOTT (FIGS 4, 50–51) Plagiotylus Scott, 1874: 272 (gen. nov. type species: Plagiotylus maculatus Scott, 1874 by monotypy); Atkinson, 1890: 125 (cat.); Reuter, 1891: 98, 160 (descr., key); Hueber, 1906: 5 (key); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 783 (cat.); Reuter, 1910: 148 (cat.); Stichel, 1933: 235 (key); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 30



631

Figure 50. Scanning electron micrograph images of Plagiotylus maculatus (male). A, lateral view; B, head and thorax, anterodorsal view; C, head and prothorax, lateral view; D, meso- and metathorax, lateral view; E, tarsus; F, pygophore, lateral view (note: most setae missing from specimen).

(cat.); Wagner & Weber, 1964: 285 (descr., key); Wagner, 1973: 100 (descr., key); Schuh, 1995: 69 (world cat.). Diagnosis: This genus is distinguished from all other Halticini by the following combination of characters: glossy surface; uniform pale green and yellow coloration; MTG obsolete; females with coleopteroid hemelytra. Redescription: Coloration (Fig. 4): mostly pale green, with yellow, brown, or black markings. Head, pronotum, and scutellum sometimes with blackish markings, antennae often with some brown or black apically, hemelytra sometimes reddened, legs generally pale. Surface and vestiture (Figs 4, 50A–F): body shining and impunctate. Body clothed in long, dark simple setae (Plagiotylus dispar with both dark and

pale setae); antennae and legs spinose, spines longest on AI and tibiae. Structure: male elongate and macropterous, female oval, coleopteroid. Head (Figs 4, 50A–C): transverse, slightly broader than anterior margin of pronotum; approximately as tall as broad, genae height greater than eye height; posterior margin of vertex straight, weakly dorsally rounded; vertex with shallow, narrow, transverse depression, posterior margin weakly upturned and carinate; frons broadly medially rounded, sulcate at clypeus. Labium (Fig. 50A, C): LI short and thick; generally reaching meso- or metacoxae. Antennae (Figs 4, 50A–C): adjacent to lower margin of eye; always shorter than body, shortest in female; AI slightly swollen, almost twice length of eye height. Thorax (Figs 4, 50A–D): pronotum trapezoidal and sloping in males, shorter, subrectangular and flat in females, collar present in males, thin and flat; callosite region weakly defined,


632


Figure 51. Male and female genitalia of Plagiotylus maculatus. A, aedeagus; B, secondary gonopore; C, left paramere; D, right paramere; E, bursa copulatrix; F, first gonapophyses, ventral view; G, posterior wall.

lateral margins sloping, posterior margin straight to weakly medially concave; mesoscutum visible in both sexes; metathoracic spiracle elongate-oval, without evaporative bodies; MTG obsolete. Hemelytra (Figs 4, 50A): macropterous male – elongate, lateral margins subparallel to apex of cuneus; clavus apically expanding; embolium narrow, R + M vein running nearly to cuneus; cuneus elongate and narrow; membrane with two cells. Coleopteroid female – clavus and R + M vein faint, cuneus and membrane absent. Legs (Figs 4, 50E): slightly longer in males; metafemora somewhat swollen in females; pretarsi without fleshy pulvilli. Abdomen (Figs 4, 50A): male: elongate, parallel-sided to pygophore. Female: broadly elongate-oval. Male genitalia (Figs 50A, F, 51A–D): pygophore conical, genital opening large; phallotheca

broad, apically tapered, with small lateral flanges, apicodorsally sulcate and weakly sclerotized; left paramere shorter than right, L-shaped, with broad sensory lobe and narrow apical apophysis, apex typically weakly hooked; right paramere much longer than left, apical club weakly laterally deflected, narrow, weakly concave, tapered at apex; ductus seminis short, thick, with flexible ribbing; secondary gonopore elongate and scoop-shaped with prised operculum, opens ventrally, apically dorsoventrally compressed, with distinct scale-like texture; endosoma simple, without spicules, in P. maculatus (and possibly others) with weakly sclerotized fields of spines. Female genitalia (Fig. 51E, F): sclerotized rings ovate, transverse, subcontiguous, weakly sclerotized, most strongly sclerotized laterally, less so towards midline,


THE HALTICINI OF THE WORLD lateral margins and adjacent portion of DLP strongly upturned; lateral upturned margins of DLP strongly sclerotized; lateral-most region of VLP joins with rami to form paired, medially projecting, sclerotized processes; posterior wall mostly membranous, with weakly sclerotized, transverse, M-shaped process; opening to vestibulum symmetrical, lateral margins weakly swollen and weakly sclerotized. Diversity and distribution: Plagiotylus includes six species, all restricted to the Mediterranean region. Included species: Plagiotylus bolivari (Reuter, 1880) Spain Plagiotylus dispar Reuter, 1899* Syria; Israel Plagiotylus maculatus Scott, 1874* Spain Plagiotylus ruffoi Tamanini, 1960 Italy Plagiotylus sahlbergi Reuter, 1901 Algeria; Greece; Libya; Tunisia Plagiotylus zorzii Tamanini, 1955 Italy Biology and host plant associations: The limited biology and host information available for Plagiotylus comes primarily from Wagner (1973). Plagiotylus maculatus is said to prefer dry, sunny areas and is found on Teucrium chamaedrys (Lamiaceae). This species has a single generation per year, overwinters as eggs, with adults present in June and July. Plagiotylus ruffoi has been collected at high altitude (1200–1890 m) in Sicily, on Astragalus siculus nebrodensis (Fabaceae), with adults are present in July. Tamanini (1960) also reports this species on an unidentified species of Prangos (Apiaceae) (Table 1). Remarks: This redescription is based on examination of P. dispar and P. maculatus, as well as the literature. Wagner (1973) provided descriptions, illustrations, and a key to all species; however only the parameres and aedeagus of Plagiotylus sahlbergi are illustrated. The relative placement of Plagiotylus differs between our equal weights and implied weights analyses, and its true relationship within the Halticini remains uncertain.

SCHOENOCORIS COSTA (FIGS 4, 52–53) Schoenocoris Reuter, 1890: 253 (gen. nov.; type species: Phytocoris flavomarginatus Costa, 1842 by monotypy); Reuter, 1891: 64, 159 (descr., key); Hueber, 1906: 4, 9 (key, descr.); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 789 (cat.); Reuter, 1910: 148 (cat.); Stichel, 1933: 235 (key); Hedicke, 1935: 59 (key); Wagner, 1952: 96, 106 (key, descr.); Carvalho,

633

1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 31 (cat.); Wagner & Weber, 1964: 275 (descr.); Wagner, 1973: 54 (descr.). Diagnosis: Recognized by the following combination of characters: head transverse, broader than posterior of pronotum; pronotum short, trapezoidal, lateral margins subparallel; hemelytra with yellow costal margins. Redescription: Small, males macropterous, females brachypterous and oblong-oval. Coloration (Fig. 4): mostly dark brown with lighter yellow and yellowbrown markings. Hemelytra with yellow costal margins. Surface and vestiture (Figs 4, 52A–H): head smooth, pronotum weakly rugulose. Body with semi-erect simple setae, erect and semi-erect spinelike setae, and adpressed scale-like setae. Head with long spine-like setae and decumbent scale-like setae. Antennae with semi-reclining spine-like setae, AI with several long spines. Pronotum and scutellum with semi-erect simple setae. Setae on thoracic pleura flat and scale-like. Hemelytra and abdomen with caudally directed simple and scale-like setae. Legs with both simple and spine-like setae, spines at apex of femora and on tibiae long and thick. Structure: head (Figs 4, 52A–D): transverse, wider than pronotum; vertex flat, posterior margin rounded, level with pronotum; frons broadly rounded; genae height approximately twice eye height; eyes large, round, bulging, substylate; buccula thin. Labium (Fig. 52D): extends to metacoxae; LI enlarged. Antennae (Figs 4, 52A–D, F): insertion below lower margin of eye; shorter than body; AI nearly twice length of eye height; AII weakly apically clavate; AIII and AIV thin. Thorax (Figs 4, 52A, B, D, E): pronotum trapezoidal, level, very short, collar faintly visible in males, callosite region bilaterally tumescent, lateral margins rounded, posterior of humeral angles rounded, posterior margin concave; mesoscutum visible in males; scutellum small; metathoracic spiracle prominent and tear-shaped, with evaporative bodies along anterior margin extending dorsally to hemelytra; MTG external efferent system rounded, extends dorsally to metathoracic spiracle, ostiole directed ventrally, peritreme above ostiole, rounded, surrounded by evaporative bodies. Hemelytra (Figs 4, 52A): macropterous male – parallelsided; cuneus elongate, narrow; membrane with two cells, extends beyond apex of abdomen. Brachypterous female – undivided; posterior margin concave, shortest at claval commissure; approximately half of abdomen exposed. Abdomen: narrow, parallel-sided in males, pear-shaped in females. Legs (Fig. 4): slightly longer in males; metafemora somewhat swollen. Male genitalia (Figs 52F–H, 53A–C): pygo-


634


Figure 52. Scanning electron micrograph images of Schoenocoris flavomarginatus (male). A, dorsum; B, head and pronotum, dorsal view; C, head, anterior view; D, head and thorax, lateral view; E, meso- and metathorax, lateral view; F, venter; G, pygophore, posterior view; H, pygophore, lateral view.

phore conical, posteroventral margin sulcate below left paramere; both parameres of similar size, with long, thin base; left paramere L-shaped, sensory lobe broad, apophysis tapering to blunt apex; right paramere extending slightly out of pygophore, apically spoon-shaped, apical margin slightly recurved and weakly, medially cleft; phallotheca elongate-oval,

tapering to blunt apex, opening weakly sclerotized on dorsal margin; ductus seminis relatively short with flexible ribbing; secondary gonopore horseshoeshaped, strongly sclerotized basally, apically becoming indistinct with faint, scale-like texturing; endosoma with two elongate, irregularly dentate, flat spicules, basally joined with small, weakly sclero-



635

Figure 53. Male and female genitalia Schoenocoris flavomarginatus. A, aedeagus; B, left paramere; C, right paramere; D, posterior wall; E, bursa copulatrix, ventral view; F, first gonapophyses, ventral view.

tized strip. Female genitalia (Fig. 52D, E): sclerotized rings small, elongate-oval, transverse, widely separated, lateral margins weakly upturned, adjacent portion of DLP sclerotized and upturned; DLP finely textured with minute spines, anteromedially cleft; VLP strongly sclerotized, forming two rectangular lobes abutting anteromedially; rami with small weakly sclerotized, medially directed lobes caudal to sclerotized rings; posterior wall of bursa copulatrix simple with weak anterolateral sclerotization, medially with bilaterally symmetrical, posteriorly tapered and diverging strips; opening to vestibulum symmetrical, with small sclerotized strips at anterior margins.

Diversity and distribution: Represented by a single species, Schoenocoris is found only in southern Europe. Included species: Schoenocoris flavomarginatus (Costa, A., 1842)* southern Europe Biology and host plant associations: Schoenocoris lives on Juncaceae, with adults present in July and August (Wagner, 1973). Remarks: Schoenocoris is most similar to Dimorphocoris and Compositocoris in general body structure. The male genitalia are also similar: in all three


636


Figure 54. Scanning electron micrograph images of Scirtetellus brevipennis. A, head and thorax, dorsal view; B, head, anterior view; C, head and prothorax, ventral view; D, head and thorax, lateral view; E, tarsus; F, pygophore, ventral view; G, pygophore, dorsal view; H, pygophore, lateral view (note: most setae missing from specimen).

genera the endosoma bears paired sclerites that are possibly medially joined and situated immediately in front of the secondary gonopore. Despite these similarities, our phylogeny does not place Schoenocoris with these genera. Additional data will likely be needed to assess accurately the phylogenetic placement of this genus.

SCIRTETELLUS REUTER (FIGS 4, 54–56) Scirtetellus Reuter, 1890: 253 (gen. nov.; type species: Labops brevipennis Reuter, 1879 by monotypy); Reuter, 1891: 67, 159 (descr., key); Hueber, 1906: 4 (key); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 789 (cat.); Reuter, 1910: 148 (cat.); Kiritshenko, 1951: 126 (key); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 66 (key); Carvalho, 1958: 31 (cat.); Kerzhner, 1964a: 964 (diag.); Medvedeva, 1975: 73 (descr., disc., key);

Schuh, 1995: 70 (world cat.); Drapolyuk & Kerzhner, 1999: 87 (descr.); Simov, 2005: 213 (descr.). Diagnosis: Recognized by the following combination of characters: both sexes short and oval, with staphylinoid hemelytra; mostly black, often with yellow or orange-yellow markings; metafemora incrassate; ductus seminis with flexible ribbing throughout; secondary gonopore heavily sclerotized, dorsoventrally compressed, opening narrow with semiclosed operculum, apically with scale-like texturing; endosoma with one to three spine- or plate-like spicules; posterior wall of bursa copulatrix sclerotized, often with prominent inter-ramal tumescences. Redescription: Both sexes staphylinoid. Coloration (Fig. 4): mostly black, almost always with yellow or orange-yellow coloration on head, sometimes with



637

Figure 55. Male and female genitalia of Scirtetellus brevipennis. A, aedeagus, phallotheca removed; B, left paramere; C, right paramere; D, posterior wall; E, bursa copulatrix, dorsal view.

yellow on pronotum and scutellum, hemelytron usually with yellow margins, abdomen sometimes with two rows of yellow spots, occasionally merged into two lines, legs brown or black. Surface and vestiture (Figs 4, 54A–H): glossy and impunctate; vertex with faint wrinkles radiating from centreline; posterior of pronotum rugulose. Body with thin simple setae; antenna with spine-like setae, AI with a few longer spines on inner surface; legs with semi-erect spines, most dense on tibiae. Structure: head (Figs 4, 54A–D): transverse, broader than pronotum, as tall as broad; posterior of vertex medially level with pronotum; eyes globular, substylate, encroaching on pronotum; genae height equal to eye height; frons broadly rounded, steeply sloping; buccula very thin; labium (Fig. 54C, D): reaches to metacoxae; LI short, moderately swollen. Antennae (Figs 4, 54A–D): insertion in line with or just below lower margin of eye; longer than body; AI swollen, slightly longer than eye height. Thorax (Figs 4, 54A, C, D): pronotum short flat, rect-

angular, collar indistinct, lateral margins rounded, callosite region indistinct to weakly defined, posterior margin straight to weakly medially cleft; mesoscutum not visible; scutellum transverse; metathoracic spiracle large and round, surrounded by evaporative bodies; MTG external efferent system evaporative area large and swollen, ostiole opens ventrolaterally, peritreme broadly tongue-shaped, orientated vertically above ostiole. Hemelyra (Figs 4, 54A): staphylinoid; short and pad-like, undivided, only covering base of abdomen. Legs: metafemora incrassate and laterally compressed; metatibiae long; pretarsi without fleshy pulvilli. Abdomen (Fig. 4): broad, parallel-sided to elongate oval, broader in females. Male genitalia (Figs 54F–H, 55A–C): pygophore conical, genital opening large, posterior margin weakly concave below left paramere; left paramere L-shaped, shorter than right, sensory lobe small but distinct, apical apophysis thin, curved, apically hooked or bifid; right paramere longer than left, api-


638


Figure 56. Female genitalia of Scirtetellus seminitens, Scirtetellus gudali, and Scirtetellus brachycerus. Scirtetellus seminitens: A, bursa copulatrix, dorsal view; B, vestibulum; C, posterior wall; D, bursa copulatrix, ventral view. Scirtetellus gudali: E, posterior wall. Scirtetellus brachycerus: F, posterior wall. Abbreviations: MP, median process formed by ventral labiate plate and ramus; VLP, ventral labiate plate.

cally concave and spoon-shaped, with minute apical apophysis; phallotheca sometimes with dense field of minute spines on outer surface near apex; ductus seminis elongate and narrow with flexible ribbing; secondary gonopore heavily sclerotized, dorsoventrally compressed, opening narrow with semi-closed operculum, apically with scale-like texturing; endosoma with one to three spine- or plate-like spicules; in some cases one spicule is enclosed in a separate pocket of the endosoma. Female genitalia (Figs 55D, E, 56A–E): sclerotized rings variable, widely separated, lateral margins sometimes weakly upturned

(e.g. Scirtetellus gudali) to strongly upturned (e.g. Scirtetellus seminitens), often with medial margin also upturned; lateral portion of DLP adjacent to sclerotized rings weakly to strongly upturned; VLP sclerotized, anteromedially divided or whole; lateralmost region of VLP joins with rami below sclerotized rings to form paired medially directed sclerotized projections, sometimes covered with fields of spines, converging with posterior margin of the posterior wall; posterior wall variable, bilaterally sclerotized in all species examined, sometimes with small, paired tumescences or small projections, in others with


THE HALTICINI OF THE WORLD elaborate inter-ramal tumescences that may be dentate and covered in dense fields of spines; opening of vestibulum symmetrical, margins weakly sclerotized and sinuous. Diversity and distribution: Scirtetellus includes 16 species in central and north Asia. Included species: Scirtetellus alashanensis Nonnaizab & Bao, 2006 Mongolia Scirtetellus bianchii Medvedeva, 1975 Asiatic Russia Scirtetellus brachycerus Kerzhner, 1962 Kazakhstan; Asiatic Russia; Mongolia Scirtetellus brevipennis (Reuter, 1879)* Turkestan Scirtetellus gudali Kiritshenko, 1951* Georgia, Russia, Asiatic Russia Scirtetellus kerzhneri Medvedeva, 1975 Asiatic Russia Scirtetellus maculiventris Kiritshenko, 1952 Asiatic Russia; Tajikistan Scirtetellus micans Medvedeva, 1975 Asiatic Russia Scirtetellus mongolicus Drapolyuk & Kerzhner, 1999 Mongolia Scirtetellus obscurus Medvedeva, 1975 Asiatic Russia Scirtetellus pallidus Medvedeva, 1975 Asiatic Russia Scirtetellus petrovi Simov, 2005* Pakistan Scirtetellus schamili Kiritshenko, 1951 Azerbaijan; Asiatic Russia Scirtetellus seminitens Horváth, 1904* Turkestan; Asiatic Russia Scirtetellus variabilis Medvedeva, 1975 Asiatic Russia Scirtetellus vittatus Kiritshenko, 1951 Russia; Asiatic Russia Biology and host plant associations: Scirtetellus is a polyphagous genus, with most species known from high mountainous regions of central Asia, in alpine and subalpine meadows from 1600 to 4800 m (Medvedeva, 1975). Species are found on various plants of the families Asteraceae, Ranunculaceae, and Poaceae (Medvedeva, 1975). Kirishenko (1952) stated that Scirtetellus are zoogeographically and ecologically most similar to Myrmecophyes (Table 1). Remarks: The most comprehensive treatment of Scirtetellus is that of Medevedva (1975, English translation 1977), which includes illustrations of male genitalia and a key to all but the most recently described species, Sc. alashanensis Nonnaizab & Bao (2006) and Sc. petrovi Simov (2005). Until now however, the female genitalia have never been docu-

639

mented. The presence of inter-ramal lobes on the posterior wall (88-0) of some species is shared with Anapus and Labops, suggesting a possible close affinity, as reflected in our phylogeny. Additionally, the presence of paired sclerotized lobes formed by fusion of the VLP and the rami (84-1) is also shared by several species of Scirtetellus and Labops, indicating that these structures may be functionally linked.

STRONGYLOCORIS BLANCHARD (FIGS 4, 57–58) Strongylocoris Blanchard, 1840: 140 (gen. nov.; type species: Strongylocoris leucocephalus Linnaeus, 1758 by monotypy); Costa, 1853: 48 (descr.); Puton, 1886: 53 (cat.); Atkinson, 1890: 119 (cat.); Reuter, 1891: 26, 157 (descr., key); Saunders, 1892: 258 (key, descr.); Hueber, 1906: 2, 32 (key, descr.); Kirkaldy, 1906: 130 (cat.); Reuter, 1910: 148 (cat.); Oshanin, 1910: 799 (cat.); Van Duzee, 1916: 211 (key); Van Duzee, 1917: 276 (cat.); Van Duzee, 1921: 135 (key spp.); Knight, 1923: 498 (key); Blatchley, 1926: 803 (key spp.); Stichel, 1933: 235 (key); Hedicke, 1935: 58 (key); Knight, 1938: 1 (note); Knight, 1941: 75, 78 (key, key spp.); Hsiao, 1942: 254 (key); China, 1943: 267 (cat.); Froeschner, 1949: 139 (key spp.); Slater, 1950: 52 (female genitalia); Kiritshenko, 1951: 127 (key); Wagner, 1952: 95, 99 (key, descr.); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 68 (key); Wagner, 1956: 277 (descr., disc. of generic limits); Carvalho, 1958: 32 (cat.); Southwood & Leston, 1959: 246 (key); Wagner, 1961: 49 (key, diag.); Kerzhner, 1964a: 967 (diag., key); Wagner & Weber, 1964: 258 (syn., descr., key); Wagner, 1973: 18 (descr., key); Schuh, 1995: 71 (cat.). Stiphrosomus Fieber, 1858: 312 [gen. (syn. by Reuter, 1888: 284); Carvalho, 1958: 33 (cat.)]. Stiphrosoma Fieber, 1861: 280 (emendation of Stiphrosomus Fieber, 1858). Diagnosis: Characterized by the following combination of characters: head broad and short, posterior margin closely adpressed to pronotum; pronotum broad, with posterior margin weakly declivent and convex; pronotum and hemelytra punctate; bursa copulatrix complex (Fig. 58G). Redescription: Broad and oval, both sexes macropterous, in some species male longer and more slender than female. Coloration (Fig. 4): variable, often black, brown, or tinged red or blue, often with yellow or yellow-brown markings. Legs sometimes concolourous, sometimes partly yellow, yellow-brown, or red. Surface and vestiture (Figs 4, 57A–G): body clothed in dark or pale simple setae; often with undercoating of short setae interspersed with longer setae; antennae and legs with semi-erect spines. Head glossy, sometimes finely rugulose; pronotum and hemelytron


640


Figure 57. Scanning electron micrograph images of Strongylocoris niger (male). A, head and pronotum, dorsal view; B, head, anterior view; C, head and thorax, lateral view; D, meso- and metathorax, lateral view; E, tarsus; F, pygophore, lateral view; G, pygophore, ventrolateral view.

densely, shallowly punctate, callosite region bilaterally impunctate; scutellum sometimes rugulose or punctuate; remainder of body impunctate. Structure: head (Figs 4, 57A–C): transverse, slightly broader than anterior of pronotum, broader than tall; vertex with transverse bisinuate depression, posterior margin slightly concave, carinate, generally touching

pronotum or nearly so; eye short and tall; head height about two times eye height; frons medially rounded, laterally shallowly depressed; base of clypeus not strongly delimited from frons; bucculae very thin. Labium (Fig. 57C): LI thick, length approximately equal to genae height. Antennae (Figs 4, 57A): insertion well below eyes; short, less than half body length;



641

Figure 58. Male and female genitalia of Strongylocoris leucocephalus. A, left paramere; B, right paramere; C, aedeagus; D, secondary gonopore; E, posterior wall; F, first gonapophyses, ventral view; G, bursa copulatrix, dorsal view.

AI not swollen, length subequal to eye height. Thorax (Figs 4, 57A–D): pronotum trapezoidal, broad, collar obsolete; callosite region weakly defined, humeral angles smoothly rounded, posterior margin weakly declivent, rounded or straight; mesoscutum not visible; scutellum small, weakly convex; metathoracic spiracle small and tear shaped, surrounded with thin strip of evaporative bodies; MTG obsolete. Hemelytra (Fig. 4): broad, lateral margin evenly rounded; R + M

vein long and close to costal margin; cuneus sort and broad; membrane short, with two cells; membrane and cuneus often steeply declivent. Legs (Figs 4, 57E): short; pretarsi without pulvilli. Male genitalia (Figs 57F, G, 58A–C): pygophore conical, basally broad; both parameres extending somewhat from pygophore; left paramere L-shaped, sensory lobe broad and rounded, apophysis curved, sometimes twisted, apically hooked; right paramere extremely


642


long, apically spoon-shaped; phallotheca short, thick, abruptly narrowing from midpoint to apex (in Strongylocoris leucocephalus apical portion compressed from all sides to form dorsal, ventral, and lateral carinas, dorsally with scale-like texturing); ductus seminis short and thick, with flexible ribbing; secondary gonopore elongate and cupped, with prised operculum opening ventrally; endosoma simple, in at least one species (St. leucocephalus) with three or four minute sclerotized spicules. Female genitalia (Fig. 57D, E): structure of bursa copulatrix complex; sclerotized rings laterally and posteriorly strongly sclerotized, becoming indistinct anteriorly, lateral and medial margins strongly upturned; DLP and VLP closely adpressed in parts, sclerotized, portions adjacent to either side of sclerotized rings folded upwards with rings, medial portion forming a curved, cylindrical enclosure; VLP adjacent to rami weakly sclerotized; DLP medially forming a weakly sclerotized plate; posterior wall simple and membranous, laterally sometimes with fields of spines; opening of vestibulum symmetrical, medial margins of first gonapophyses moderately swollen, medial margins of vestibular opening weakly sclerotized.

Diversity and distribution: Strongylocoris exhibits a Palaearctic distribution, with most of the 16 species found in the Mediterranean.

Included species: Strongylocoris amabilis (Douglas & Scott, 1868) Egypt; Israel Strongylocoris atrocoeruleus (Fieber, 1864) south Europe Strongylocoris cicadifrons Costa, 1853 Europe; Israel Strongylocoris coerulescens Lindberg, 1940 Morocco Strongylocoris enki Linnavuori, 1984 Iraq Strongylocoris erythroleptus Costa, 1853 Italy Strongylocoris ferreri Ribes & Pagola-Carte, 2007 Spain Strongylocoris franzi Wagner, 1955 Spain Strongylocoris leucocephalus (Linnaeus, 1758)* Palaearctic Strongylocoris luridus (Fallén, 1807)* Europe Strongylocoris niger (Herrich-Schäeffer, 1835) Europe Strongylocoris oberthuri Reuter, 1905c France Strongylocoris obscurus (Rambur, 1839) Mediterranean Strongylocoris raimondoi Carapezza, 1991 Italy Strongylocoris seabrai Schmidt, 1939 Portugal; Spain Strongylocoris steganoides (Sahlberg, 1875)* Austria; Russia; Sweden

Biology and host plant associations: Host species are recorded for nine species of Strongylocoris (Table 1). Virtually all are associated with asterids, whereas a single species, St. luridus, has been collected on the rosid species Calicotome villosa (Fabaceae) (Pericart, 1965). Remarks: Three species of Strongylocoris were dissected in this study (St. leucocephalus, St. luridus, and St. steganoides). In all three the female genitalia bear the same diagnostic characters, most notably the unique structure of the bursa copulatrix. More specimens must be examined in order to determine the degree of variation in this character complex across the genus.

DISCUSSION The Halticini is retained within the subfamily Orthotylinae on the basis of the apically convergent lamellate parempodia. This pretarsal character state also occurs in the Phylinae: Pilophorini, but the Halticini lack the phyline-type male genitalia (s-shaped endosoma, phallotheca attached directly to the pygophore). In contrast, the halticine genitalia are similar to those found in the nominotypical tribe Orthotylini, with robust, strap-like spicules and an oval secondary gonopore. The Halticini have previously been defined by the absence of inter-ramal lobes (= K-structures sensu Slater, 1950); however, we found a great diversity in the morphology of the posterior wall of the bursa copulatrix, including the presence of inter-ramal lobes in three halticine genera, which we believe are homologous to those found in the Orthotylini and the Austromirini Carvalho. An absence of inter-ramal lobes is the norm for most Halticini, and is also observed in the Nichomachini, and the newly erected monotypic tribe Coridromiini. On the basis of previous definitions of the Halticini, these observations render the tribe as lacking defining synapomorphies. Schuh & Lattin (1980), in their paper on the first Nearctic Myrmecophyes species, made the observation that this species, M. oregonensis, and another halticine, Anapus americanus, possessed jointly symmetrical and non-overlapping sclerotization of the opening of the vestibulum associated with the first gonapophyses; they concluded that this character state was worthy of broader comparative investigation in terms of establishing an alternative synapomorphy for the tribe. Subsequently, Pluot-Sigwalt & Matocq (2006) found similar symmetry for additional halticine species. We too have found such symmetry in the vestibulum of all halticine females examined; however such symmetry is also present in the Nichomachini, the Coridromiini (= Coridromius), and


THE HALTICINI OF THE WORLD several Orthotylini, such as the Hadronema group (Forero, 2008) and Harveycapsus dimorpha (Cassis, Symonds & Tatarnic, 2010), and can thus not be considered as unique to the tribe. From the present study a single synapomorphy has been identified: the presence of elongate left and right paramere bases. This is not unusual in the speciose family Miridae, where great morphological diversity exists, and higher taxonomic level synapomorphies are rare. Our definition of the Halticini retains the tribe as a highly diverse suprageneric group with a number of highly autapomorphic genera (e.g. Chorosomella, Dicyphopsis). This study has produced robust definitions of the genera, inclusive of information for the male and female genitalia, with exemplars illustrated for nearly all genera. On the basis of external characters such as head structure, wing polymorphism, the external efferent system of the MTG, and the genitalia, we have mostly supported the previous generic status of the halticines as summarized in the world catalogue of Schuh (1995). Apart from the removal of Coridromius from the Halticini, the most significant changes we present are in the nominate genus Halticus, which as previously defined was not monophyletic. Our subdivision of the genus leads to the following outcomes: (1) Halticus has been redefined to include only those species, which as a group have a Holarctic distribution, and are defined by large, transverse, subcontiguous, female sclerotized rings, the scoop-shaped secondary gonopore, and vestigial endosoma; (2) Ectmetopterus is redefined to include east Asian species previously placed within Halticus and now recognized by the tripartite left paramere; and (3) Microtechnites has been removed from synonymy, with its generic status restored for a group of mostly Neotropical species defined by the multiple spicules and the small and widely separated female sclerotized rings. Based on our phylogenetic analysis the present evidence does not support any particularly close affinity amongst these three genera. In summary, this study is the first global assessment of halticines, including the comparative examination of existing and new character systems. The discovery in some genera of inter-ramal tumescences of the posterior wall putatively homologous to inter-ramal lobes of the Orthotylini refutes previous definitions of the Halticini; however the confirmation that in all genera the margins of vestibular opening are symmetrical provides an additional (although homoplasious) defining trait for the tribe. Subsequent studies of other mirids should involve examination of this character system to determine its usefulness in higher-level taxonomic delineation. This study will inform future phylogenetic treatments of the Halticini, by providing useful, detailed character systems. The lack of strong

643

phylogenetic support for all but a few clades suggests that future studies may benefit from the inclusion of molecular data, in order to ascertain accurately the relationships of highly autapomorphic genera such as Chorosomella, for which morphological character homology is difficult to confirm.

ACKNOWLEDGEMENTS We wish to thank the museums mentioned herein for supplying the material for this work. We are also grateful to the following individuals for providing fresh material for study: freshly collected specimens of Myrmecophyes alboornatus were provided by Fedor Konstantinov of the Zoological Institute of Saint Petersburg; specimens of Halticus apterus were provided by Michael D. Schwartz of the Agriculture and Agri-Food Canada (CNC, Ottawa); Randall T. Schuh from the American Museum of Natural History provided specimens of Compositocoris senecionus; Thomas Henry from the U.S. Natural History Museum and Dimitri Forero (AMNH) provided specimens of recent collections of Cafayatina altigena from Argentina. Additionally, Nikolay Simov provided specimens of Dimorphocoris fuscus from Bulgaria, and the recently described Scirtetellus petrovi from Pakistan. We also thank Sue Lindsay (Australian Museum) for providing assistance with the scanning electron micrographs of most taxa. This manuscript was significantly improved through comments from two anonymous reviewers. Lastly, we thank Michael Schwartz, who provided the senior author with tremendous assistance throughout this project. This research was conducted during the course of the senior author’s PhD, which was funded through the NSF Planetary Biodiversity Inventories grant DEB0316495 to R. T. Schuh and G. Cassis.

REFERENCES Alayo DP. 1974. Los Hemipteros de Cuba. Parte XIII. Familia Miridae. Torreia 32: 1–41. Amyot CJB, Serville A. 1843. Histoire Naturelle des Insectes. Hémiptères. Paris: Fain et Thunot. Asquith A. 1995. Evolution of the endemic Hawaiian plant bug genus Sarona: speciation on geographical and ecological islands. In: Wagner WL, Funk V, eds. Speciation and biogeography of the Hawaiian biota. Washington, DC: Smithsonian Institution Press, 90–119. Atkinson ET. 1890. Catalogue of the Insecta. Order Rhynchota. Suborder Hemiptera-Heteroptera. Family Capsidae. Journal of the Asiatic Society of Bengal (Natural Science Supplement) 58: 25–200. Berg C. 1883. Addenda et emendanda ad Hemiptera Argentina. Anales de la Sociedad Cientifica Argentina 16: 5–32, 73–87, 105–125, 180–191, 231–241, 285–294.


644


Bergevin ED. 1916. Note sur le genre Dimorphocoris Reut. Bulletin de la Société d’Histoire Naturelle, Algérie 7: 61–62. Blanchard E. 1840. Hémiptères. In: de Castelnau FLL, ed. Histoire Naturelle des Animaux Articulés, Annelides, Crustaces, Arachnides, Myriapodes et Insectes, TOM III. Orthoptères, Neuroptères, Hémiptères, Hymenoptères, Lépidoptères et Diptères. Paris: P. Duméril, 1–671. Blatchley WS. 1926. Heteroptera or True Bugs of Eastern North America, with especial reference to the faunas of Indiana and Florida. Indianapolis: Nature Publishing Company. Brullé MA. 1836. Histoire Naturelle des Insectes (Coléoptères, Orthoptères et Hémiptères), traitant de leur organization et de leurs moeurs en général. Paris: F.D. Pillot. Burmeister HCC. 1835. Handbuch der Entomologie. Zweiter Band. Erste Abtheilung-Rhynchota. Berlin: Theodore Christian Friedrich Enslin. Butler EA. 1923. A biology of the British Hemiptera Heteroptera. London: H.F. and G. Witherby. Bykov AA. 1969. New species of capsid bugs from Kirgizia. Izvestiya Akademii Nauk Kirgizskoi SSR 1969: 54–55. Bykov AA. 1970. A new species of bug, Myrmecophyes piceus Bykov, sp. n. (Hemiptera, Miridae) from Kirgizia. Entomologitshskie issledovaniya v Kirgizii. 1970: 54–56. Bykov AA. 1971. Mirid bugs of the genus Myrmecophyes Fieb. (Heteroptera, Miridae) in the fauna of the Tien-Shan and the Pamir-Alai. Entomological Review 50: 495–501. Carapezza A. 1991. Three new species of Miridae collected in Sicily (Insecta, Heteroptera). Naturalista Siciliano 15: 179– 188. Carapezza A. 2002. Dimorphocoris (s. str.) poggii n. sp., a new Miridae from the Ligurian Alps (Heteroptera Miridae). Annali del Museo Civico di Storia Naturale ‘Giacomo Doria’ 94: 473–483. Carpinera J. 2001. Handbook of vegetable pests. San Diego, CA: Academic Press. Carvalho JCM. 1952. On the major classification of the Miridae (Hemiptera). (With keys to subfamilies and tribes and a catalogue of the world genera). Anais da Academia Brasileira de Ciencias 24: 31–110. Carvalho JCM. 1955. Keys to the genera of Miridae of the world (Hemiptera). Bolletim do Museu Paraense Emilio Goeldi 11: 5–151. Carvalho JCM. 1956. Insects of Micronesia. Heteroptera: Miridae. Insects of Micronesia 7: 1–100. Carvalho JCM. 1958. Catálogo dos Mirídeos do Mundo. Parte III. Subfamília Orthotylinae. Arquivos do Museo Nacional, Rio de Janeiro 47: 161. Carvalho JCM. 1959. A catalogue of the Miridae of the world. Part IV. Mirinae. Arquivos do Museo Nacional, Rio de Janeiro 48: 1–384. Carvalho JCM. 1987. The genus Coridromius Signoret, with descriptions of new species (Hemiptera, Miridae). Revista Brasileira de Entomologia 31: 61–69. Carvalho JCM, Carpintero DL. 1986. Mirideos Neotropicais; CCLXXIII: Descricoes de um novo genero e treze especies novas da republica Argentina (Hemiptera). Revista Brasileira de Biologia 46: 607–625.

Carvalho JCM, Carpintero DL. 1992. Miridos neotropicales, CCCXXXVII. Description de doce espécies nuevas de Orthotylinae (Hemiptera). Revista Brasileira de Biologia 51: 763–780. Carvalho JCM, Leston D. 1952. The classification of the British Miridae (Hem.), with keys to the genera. Entomologist’s Monthly Magazine 88: 231–251. Cassis G. 1986. A systematic study of the subfamily Dicyphinae (Heteroptera: Miridae). PhD Dissertation. Oregon State University, Corvallis. Cassis G. 2008. The Lattinova complex of austromirine plant bugs (Hemiptera: Heteroptera: Orthotylinae). Proceedings of the Entomological Society of Washington 110: 845– 939. Cassis G, Gross GF. 1995. Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). In: Houston WWK, Maynard GV, eds. Zoological catalogue of Australia. 27.3A. Melbourne: CSIRO. Cassis G, Moulds T. 2002. A systematic revision of the plantbug genus Kirkaldyella Poppius (Heteroptera: Miridae: Orthotylinae: Austromirini). Insect Systematics and Evolution 33: 53–90. Cassis G, Schwartz MD, Moulds T. 2003. Systematics and new taxa of the Vannius complex (Hemiptera: Miridae: Cylapinae) from the Australian Region. Memoirs of Queensland Museum 49: 125–143. Cassis G, Symonds S, Tatarnic N. 2010. A remarkable new species of stone-dwelling Orthotylini (Heteroptera: Miridae: Orthotylinae) from Australia. Zootaxa 2485: 58–68. Chérot F, Konstantinov F, Yasunaga T. 2004. Two new plant bug species of the genus Coridromius Signoret, 1862 from the Oriental region with a new synonymy (Heteroptera, Miridae, Orthotylinae, Halticini). Belgian Journal of Entomology 6: 57–67. China WE. 1943. The generic names of British insects. Part 8. The generic names of the British Hemiptera-Heteroptera, with a check list of the British species. London: Royal Entomological Society of London. Costa A. 1842. Mèmoire pour servir à l’histoire des Hémiptères Héteroptères de Deux-Siciles. Annales de la Société Entomologique de France 10: 279–308. Costa A. 1853. Cimicum Regni Neapolitani Centuria tertia et quartae fragmentum. Atti del Reale Istituto d’Incorragiamento alle Scienze Naturali 8: 3–77. Davis NT. 1955. Morphology of the female organs of reproduction in the Miridae (Hemiptera). Annals of the Entomological Society of America 48: 132–150. Distant WL. 1893. Insecta. Rhynchota. HemipteraHeteroptera. Biologia Centrali Ammericana, Suppl. 1-xx: 329–462. Distant WL. 1904. The fauna of British India, including Ceylon and Burma. Rhynchota, vol. II, Part 2. London: Taylor and Francis, 243–503. Distant WL. 1910a. Descriptions of Oriental Capsidae. Annual Magazine of Natural History 5: 10–22. Distant WL. 1910b. The fauna of British India, including Ceylon and Burma. Rhynchota Vol. V. Heteroptera: Appendix. London: Taylor and Francis.


THE HALTICINI OF THE WORLD Distant WL. 1914. Rhynchota from New Caledonia and the surrounding islands. Nova Caledonia Zoologie 1: 369– 390. Douglas JW, Scott J. 1865. The British Hemiptera. Vol. I. Hemiptera-Heteroptera. London: Robert Hardwicke. Douglas JW, Scott J. 1868. List of captures of Hemiptera in Palestine and Syria; together with descriptions of several new species. Entomologist’s Monthly Magazine 5: 114–118, 135–139. Drapolyuk IS. 1989. Species of the genus Myrmecophyes Fieb. (Heteroptera: Miridae) from the Caucasus. Entomologicheskoe Obozrenie 18: 125–135. Drapolyuk IS, Kerzhner IM. 1999. Species of the genus Scirtetelus from Mongolia (Heteroptera: Miridae). Zoosystematica Rossica 8: 87–88. Drapolyuk IS, Kerzhner IM. 2000. New species of Orthocephalus and Myrmecophyes from Kazakhstan, Uzbekistan and Turkmenistan (Heteroptera: Miridae). Zoosystematica Rossica 8: 301–305. Ehanno B. 1960. Contribution à la connaissance des Insectes Héteroptères Miridae Armoricains (2e note). Bulletin de la Société Scientifique de Bretagne 35: 313–324. Ehanno B. 1965. Notes écologiques sur les Miridae (InsectaHétèroptera) observés en Bretagne sur le chene. Vie et Milieu 16: 517–533. Ehanno B. 1991. Contribution à l’étude des Miridae, compléments à la description de Myrmecophyes alboornatus (Stal, 1858) (Halticinae, Halticini, Myrmecopharia). Bulletin de la Société Scientifique de Bretagne 62: 25–36. Ehanno B. 1992. Compléments à la faune de France (et pays voisins) des Hétéroptères Miridae Hahn, 1831. V – des espèces du genre Dimorphocoris Reuter, 1891 (Halticinae, Halticini, Laboparia) 1 – espèces des Alpes (France, Autriche, Italie), des Apennins (Italie) et du domaine Meditérranéen: additions aux donées acquises et descriptions d’espèces nouvelles. Bulletin de la Société Scientifique de Bretagne 63: 5–187. Ehanno B. 1994a. Compléments à la faune de France et pays voisins: Espagne (Catalogne), Andorre, des Hétéroptères Miridae Hahn, 1831 – V:2 – 7. Des espèces du genre Dimorphocoris Reuter, 1891 (Halticinae, Halticini, Laboparia). Bulletin de la Société Scientifique de Bretagne 64: 5–48. Ehanno B. 1994b. Compléments à la faune de France et pays voisins: Espagne (Catalogne), Andorre, des Hétéroptères Miridae Hahn, 1831. (5 – 7. Additions aux donées acquises: 5 – 1. (Dim. 1, 1992)). Bulletin de la Société Scientifique de Bretagne 64: 203–208. Ehanno B, Ribes J. 1994. Compléments à la faune de France et de pays voisins des Hétéroptères Miridae Hahn, 1831. V -des espèces du genre Dimorphocoris Reuter, 1891 (Halticinae, Halticini, Laboparia) – 5 – Espèces des Pyrenées (France, Espagne, Andorre). Additions aux donées acquises et descriptions d’espèces nouvelles. Bulletin de la Société Scientifique de Bretagne 64: 49–202. Fabricius JC. 1777. Genera Insectorum eorumque characteres naturales secundum numerum figuram situm et proportionem omnium partium oris adjecta mantissa specierum nuper detectarum. Chilonii: Mich. Friedr. Bartschii.

645

Fallén CF. 1807. Monographia cimicum sveciae. Hafniae: C. G. Proft. Fallén CF. 1829. Hemiptera Sueciae. Cimicides eorumque familae affines. London. Fieber FX. 1858. Criterien zur generischen Theilung der Phytocoriden (Capsini auct.). Wiener Entomologische Monatschrift 2: 289–327, 329–347, 388. Fieber FX. 1860a. Die europäischen Hemipteren Halbflügler (Rhynchota Heteroptera). Nach der analytischen Methode bearbeitet. Part I. Wien: C. Gerold. Fieber FX. 1860b. Exegesen in Hemiptera. Wiener Entomologische Monatschrift 4: 257–272. Fieber FX. 1861. Die europäischen Hemiptera Halbflügler (Rhynchota Heteroptera). Nach der analytischen Methode bearbeitet. Part II. Wien: C. Gerold. Fieber FX. 1864. Neuere Entdeckungen in europäischen Hemipteren. Wiener Entomologische Monatschrift 8: 65–86, 205–236, 321–336. Fieber FX. 1870. Dodecas neuer Gattungen und neuer Arten europäischer Hemiptera. Verhandlungen der ZoologischBotanischen Gesellschaft in Wien 20: 243–264, pls. 5–6. Flor G. 1860. Die Rhynchoten Livlands in Systematischer Folge bescreiben. Erster Theil: Rhynchota frontirostria Zett. (Hemiptera Heteroptera Ast.). Archiv für die Naturkunde Liv-, Ehst-, und Kurlands. II Serie, Biologische Naturkunde (1860–1861). Dorpat: Carl Schulz. Forero D. 2008. Revision and phylogenetic analysis of the Hadronema group (Miridae: Orthotylinae: Orthotylini), with descriptions of new genera and new species, and comments on the Neotropical genus Tupimiris. Bulletin of the American Museum of Natural History 312: 1–172. Franz H, Wagner E. 1961. Die Nordost-Alpen im Spiegel ihrer Landtierwelt, II. Innsbruck: Universitatsverlag Wagner. Froeschner RC. 1949. Contributions to a synopsis of the Hemiptera of Missouri, Pt IV. Hebridae, Mesoveliidae, Cimicidae, Anthocoridae, Cryptostemmatidae, Isometopidae, Miridae. American Midland Naturalist 42: 123–188. Fuente JM. 1918. Nota hemipterologica. Boletin de la Sociedad Aragonesa de Ciencias Naturales 17: 236–239. Fuxa JR, Kamm JA. 1976. Dispersal of Labops hesperius on Rangeland. Annals of the Entomological Society of America 69: 891–893. Geoffroy EL. 1785. [Description of new taxa]. In: de Fourcroy AF, ed. Entomologia Parisiensis; sive catalogus insectorum quae in Agro Parisiensi reperiuntur; secundum methodum Geoffroeanam in sectiones, genera et species distributus; cui addita sunt nomina trivialia et fere trecentae novae species. Paris: Via et Aedibus Serpentineis. Göllner-Scheiding U. 1970. Beiträge zur HeteropterenFauna Brandenburgs. 1. Die Heteropteren-Fauna des Gross-Machnower Weinbergs und seiner naheren Umgebung. Archiv fuer Naturschutz und Landschaftsforschung 10: 41–70. Göllner-Scheiding U. 1972. Beiträge zur HeteropterenFauna Brandenburgs. 2. Übersicht über die Heteropteren von Brandenburg. Veroffentlichungen Bizirksheimat Mus. Potsdam 25/26: 5–39.


646


Goloboff PA. 1993. Estimating character weights during tree search. Cladistics 9: 83–91. Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786. Gravestein WH. 1978. Hemiptera Heteroptera new to the Baleares, in particular to the island of Mallorca. Entomologische Berichte 38: 37–39. Hahn CW. 1833. Die wanzenartigen insecten, vol. I. Nürnburg: In der C.H. Zeh’schen Buchhandlung. Hahn CW. 1835. Die wanzenartigen insecten. Vols. II & III. Nürnberg: In der C.H. Zeh’schen Buchhandlung. Hedicke H. 1935. Heteroptera. Die Tierwelt Mitteleuropas (IV) 3: 15–70. Henry T. 1983. The garden fleahopper genus Halticus (Hemiptera: Miridae): ressurection of an old name and key to species of the Western Hemisphere. Proceedings of the Entomological Society of Washington 85: 607–611. Henry T, Smith CL. 1979. An annotated list of the Miridae of Georgia (Hemiptera-Heteroptera). Journal of the Georgia Entomological Society 14: 212–220. Henry TJ, Wilson MR. 2004. First records of eleven true bugs (Hemiptera: Heteroptera) from the Galapagos Islands, with miscellaneous notes and corrections to published reports. Journal of the New York Entomological Society 112: 75–86. Herrich-Schäeffer GAW. 1835. Nomenclatur Entomologicus. Verzeichniss der europäischen Insecten; zur Erleichterung des Tauschverkehrs mit Preisen versehen. I. Lepidoptera & Hemiptera. Regensburg: Friedrich Prestet. Herrich-Schäeffer GAW. 1850. Die wanzenartigen insekten, vol. XIV. Nürnberg: C. H. Zeh’schen Buchhandlung. Hoberlandt L. 1956. Results of the Zoological Scientific Expedition of the National Museum in Praha to Turkey. 18. Hemiptera IV. Terrestrial Hemiptera-Heteroptera of Turkey. Acta Entomologica Musei Nationalis Pragae, Supplementum 3: 1–264. Hoberlandt L. 1963. Distributional notes on some species of Heteroptera from Czechoslovakia with a contribution on the taxonomy of the genus Hyoidea (Heteroptera, Miridae). Acta Faunistica Entomologica Musei Nationalis Pragae 9: 253– 283. Horváth G. 1877. Piezocranum, novum genus Capsidarum descripsit. Természetrajzi Füzetek 1: 92–93. Horváth G. 1880. Hemipterologiai Közlemények. Termeszetrajzi Füzetek 4: 184–192. Horváth G. 1881. Hemiptera nova vel minus cognita. Természetrajzi Füzetek 5: 39–43. Horváth G. 1884. Diagnoses Hemipterorum. Termeszetrajzi Füzetek 8: 9–15. 315–320. Horváth G. 1889. Beitrag zur Hemipteren-Fauna von Turkmenien. Wiener Entomologische Ztg 1: 169–174. Horváth G. 1898. Capsidae novae aut minus cognitae. Természetrajzi Füzetek 21: 501–505. Horváth G. 1904. Insecta Heptapotamica a D. D. Almasy et Stummer-Traunfels collecta. I. Hemiptera. Annales Historico-Naturales Musei Nationalis Hungarici 2: 574–590. Horváth G. 1905. Hémiptères nouveaux de Japon. Annales Historico-Naturales Musei Nationalis Hungarici 3: 413–423.

Horváth G. 1906. Ergebnisse einer naturwissenchaftlichen Reise zum Erdschias-Dagh (Kleinasien). Hemipteren. Annalen des K.K. Naturhistorischen Hofmuseums Wien 20: 179–189. Horváth G. 1926. Species Miridarum generis Myrmecophyes Fieber. Annales Historico-Naturales Musei Nationalis Hungarici 23: 187–197. Horváth G. 1927. Species Miridarum generis Myrmecophyes Fieber. Annales Historico-Naturales Musei Nationalis Hungarici 23: 187–197. Hsiao TY. 1942. A list of Chinese Miridae (Hemiptera) with keys to subfamilies, tribes, genera and species. Iowa State College Journal of Science 16: 241–269. Hueber T. 1906. Synopsis der deutschen Blindwanzen (Hemiptera Heteroptera Fam. Capsidae). Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg 2: 201–262. Jakovlev BE. 1881. Contributions to the fauna of the Heteropterous insects of Russia and neighbouring countries. V–VIII. Bulletin de la Société Impériale des Naturalistes de Moscou 56: 194–214. Jakovlev BE. 1875. Hemiptera Heteroptera of the Astrakhan Region. Bulletin de la Société Impériale des Naturalistes de Moscou 49: 145–174. Jakovlev BE. 1877. New bugs, Hemiptera Heteroptera of the Astrakhan fauna. Second Supplement. Bulletin de la Société Impériale des Naturalistes de Moscou 52: 269–300. Joakimov D. 1909. On the fauna of Hemiptera of Bulgaria. Sbornik za Narodni Umotvoreniya Nauka i Knizhnina 25: 1–34. Josifov M, Kerzhner IM. 1972. Heteroptera aus Korea. I Teil (Ochteridae, Gerridae, Saldidae, Nabidae, Anthocoridae, Miridae, Tingidae, und Reduviidae). Annales Zoologici, Warsaw 29: 147–180. Josifov M. 1974. Die Heteropteren der Bulgarischen Schwarzmeerkuste. Izvestiya na Zoologicheskiya Institut s Muzei 39: 5–27. Kelton LA. 1955. Genera and subgenera of the Lygus complex (Hemiptera: Miridae). Canadian Entomologist 87: 277–301. Kelton LA. 1959. Male genitalia as taxonomic characters in the Miridae. Canadian Entomologist 91: 1–71. Kelton LA. 1980. The plant bugs of the Prairie Provinces of Canada. Heteroptera: Miridae. The insects and arachnids of Canada. Part 8. Hull, Quebec: Canadian Government Publishing Centre. Kerzhner IM. 1962. Materials on the taxonomy of capsid bugs (Hemiptera – Miridae) in the fauna of the USSR. Entomologicheskoe Obozrenie 41: 372–387. Kerzhner IM. 1964. Family Isometopidae. Family Miridae (Capsidae). In: Opredelitel’nasekomykh evropeiskoichasti SSSR (Ed. by Bei-Bienko, GY), pp. 700–765. Kerzhner IM. 1964a. New and little known Heteroptera from Kazakhstan and some other regions of USSR. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 34: 113–130. Kerzhner IM. 1964b. Family Isometopidae. Family Miridae (Capsidae). In: Bei-Bienko GY, ed. Opredelitel’nasekomykh evropeiskoichasti SSSR [Keys to the insects of the European


THE HALTICINI OF THE WORLD part of the USSR], vol. 1. Apterygota, Paleoptera, Hemmimetabol, Moskva, Leningrad: Nauka, 700–765. Kerzhner IM. 1970. New and little known mirid bugs (Heteroptera, Miridae) from the USSR and Mongolia. Entomologicheskoe Obozrenie 49: 392–399. Kerzhner IM. 1978. Bugs (Heteroptera) of Saghalien and Kurile Islands. Trudy Biologo-Pochvennogo Instituta Dal’nevostostochnoe Otdelenie Akademiya Nauk SSSR, new ser 50: 31–57. Kerzhner IM. 1988. Novye i maloizvestnye poluzhestkiokrylye nasekomye (Heteroptera) Dal’nego Vostoka SSSR (1987). Akademiya Nauk USSR, Vladivostok 1987: 1–84. Kerzhner IM. 1996. On type specimens of some Palaearctic Miridae in the Hungarian Museum of Natural History (Heteroptera). Zoosystematica Rossica 5: 99–102. Kerzhner IM, Josifov M. 1999. Cimicomorpha 2. Miridae. In: Aukema B, Rieger C, eds. Catalogue of the Heteroptera of the Palaearctic Region, vol. 3. Amsterdam: Netherlands Entomological Society. Kerzhner IM, Konstantinov FV. 1999. Structure of the aedeagus in Miridae (Heteroptera) and its bearing to suprageneric classification. Acta Societatis Zoologicae Bohemicae 63: 117–137. Kerzhner IM, Muminov NN. 1974. New and little-known capsid bugs (Heteroptera, Miridae) from Middle Asia. Izvestiya Akademii Nauk Tadzhikskoi SSR 56: 42–46. Kerzhner IM, Schuh RT. 1995. Homonymy, synonymy, and new combinations in the Miridae (Heteroptera). American Museum Novitates 3137: 1–11. Kerzhner IM, Yachevskii TL. 1967. Order Hemiptera (Heteroptera). In: Bei-Bienko GY, ed. Keys to the insects of the European USSR. Vol 1. Apterygota, Paleoptera, Hemimetabola. Moskva, Leningrad: Zoological Institute, USSR Academy of Sciences, 851–1118. Kiritshenko AN. 1911. Neue und noch wenig bekante Hemipteren der russischen Fauna. Revue Russe d’Entomologie 11: 80–95. Kiritshenko AN. 1951. [True bugs of the European parts of the USSR (Hemiptera): Key and bibliography]. Opredeliteli po Faune SSSR 42: 1–423. Kiritshenko AN. 1952. [New and little known bugs (Hemiptera-Heteroptera) of Tadjikistan]. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 10: 140–198. Kirkaldy GW. 1902. Hemiptera. Fauna Hawaiiensis 3: 93–174. Kirkaldy GW. 1904. Some new Oahuan (Hawaiian) Hemiptera. Entomologist 37: 174–179. Kirkaldy GW. 1906. List of the genera of the pagiopodous Hemiptera-Heteroptera, with their type species from 1758 to 1904 and also of the aquatic and semi-aquatic Trochalopoda. Transactions of the American Entomological Society 32: 117–156, 156a–156b. Kiyak S. 1995. Replacement name for the genus Halticidea Reuter, 1901 and the subfamily Halticinae Kirkaldy, 1902 of Miridae (Insecta, Hemiptera). Beitraege Zur Entomologie 45: 215–216. Knight HH. 1922. The North American species of Labops (Heteroptera-Miridae). Canadian Entomologist 54: 258–261.

647

Knight HH. 1923. Guide to the insects of Connecticut. Part IV. The Hemiptera or sucking insects of Connecticut – family Miridae (Capsidae). State of Connecticut Geological and Natural History Survey, Bulletin 34: 422–658. Knight HH. 1929. Labops verae, new species, with Labopella, Nicholia, and Pronotocrepis, new genera of North American Miridae (Hemiptera). Canadian Entomologist 61: 214–218. Knight HH. 1938. Idatiella China: two new species, with a key (Hemiptera: Miridae). Annals and Magazine of Natural History 1: 25–27. Knight HH. 1941. The plant bugs, or Miridae of Illinois. Bulletin of the Illinois Natural History Survey 22: 1–234. Knight HH. 1959. New genera and species of North American Miridae (Hemiptera). Iowa State College Journal of Science 33: 421–426. Knight HH. 1968. Taxonomic review: Miridae of the Nevada test site and the western United States. Brigham Young University Science Bulletin, Biological Series 9. Kulik SA. 1965. New species of capsid-bugs (Heteroptera, Miridae) from East Siberia and from the Far East. Zoologicheskii Zhurnal 44: 1497–1505. Kullenberg B. 1947. Der Kopulationsapparat der Insekten aus Phylogenetischen Gesicktspunkt. Zoologische Bijdragen 25: 79–90. Laporte FL. 1832. Essai d’une classification systématique de l’ordre des Hémiptères (Hémiptères-Hétéroptères Latr.). Magasin de Zoologie 2: 1–16. Latreille PA. 1829. Les Crutacés, les Arachnides et les Insectes distribués in familles naturelles. In: Cuviere G, ed. Le Règne Animal Distribué d’après Organisation, pour Servir de Base à l’Histoire Naturelle des Animaux et d’Introduction à l’Anatomie Comparée, vol. V. Paris: Déterville, 556. Leach WE. 1819. The entomologist’s useful compendium; or an introduction to the knowledge of British insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; together with the genera of Linné, and the modern method of arranging the classes Crustacea, Myriapoda, spiders, mites, and insects from their affinities and structure, according to the views of Dr. Leach. Also an explanation of the terms used in entomology; a calendar of the times of appearance, and usual situations of near 3000 species of British insects; with instructions for collecting and fitting up objects for the microscope. London: Thomas Boys. Lindberg H. 1940. Inventa entomologica itineris Hispanici et Maroccani, quod a. 1926 fecerunt Harald et Hakan Lindberg XXVI. Die Capsidenfauna von Marokko. Societatis Scientiarium Fennica, Commentationes Biologicae 7: 1–55. Lindberg H. 1956. Über einige Miriden in E. de Bergevins Sammlung. Notulae Entomologicae 36: 53–64. Linnaeus C. 1758. Systema naturae, 10th edn, vol. 1. Salvii L, Holmiae. 823 + III pp. Linnavuori R. 1975. Hemiptera of the Sudan, with remarks on some species of the adjacent countries. 1. Miridae and Isometopidae. Annales Zoologici Fennici 12: 1–118. Linnavuori R. 1986. Heteroptera of Saudi Arabia. Fauna of Saudi Arabia 8: 31–197.


648


Linnavuori RE. 1973. A collection of Heteroptera from Katanga, with remarks on some species from other parts of the Ethiopian Region. Annales Entomologici Fennici 39: 70–94. Linnavuori RE. 1984. New species of Hemiptera Heteroptera from Iraq and adjacent countries. Acta Entomologica Fennica 44: 1–59. Linnavuori RE. 1992. The lateralis group of the genus Dimorphocoris Reuter of Greece and the Middle East (Heteroptera, Miridae, Halticini). Entomologica Fennica 3: 215– 222. Linnavuori RE. 1994. Orthotylinae of West, Central and North-East Africa (Heteroptera, Miridae). Acta Zoologica Fennica 193: 1–84. Linnavuori RE. 1997. Taxonomic studies on the Miridae (Heteroptera) of Yemen and Iran. Acta Universitatis Carolinae, Biologica 40: 301–320. Liu G-Q, Zhao RJ. 1999. New species of the genus Coridromius Signoret from China (Heteroptera: Mirinae). Acta Zootaxonomica Sinica 24: 55–58. Maldonado CJ. 1969. The Miridae of Puerto Rico (Insecta, Hemiptera). University of Puerto Rico Agricultural Experiment Station Technical Paper 45: 1–133. Medvedeva ND. 1975. Revision of Scirtetellus Reut. (Heteroptera, Miridae). Entomologicheskoe Obozrenie 54: 814– 826. [English translation: 1977. Entomological Review 54: 73–82]. Meyer-Dür LR. 1843. Verzeichnis der in der Schweiz einhimischen Rhynchoten (Hemiptera Linn.). Erstes Heft. Die Familie der Capsini. Solothurn: Jent und Gassmann. Mills HB. 1939. Montana insect pests for 1937 and 1938. Bulletin of the Montana Agricultural Experiment Station 366: 1–32. Mills HB. 1941. Montana insect pests for 1939 and 1940. Bulletin of the Montana Agricultural Experiment Station 384: 1–27. Miyamoto S, Yasunaga T. 1999. Discovery of the genus Coridromius Signoret (Heteroptera: Miridae) from Japan, with descriptions of two new species. Biogeography 1: 33–37. Montrouzier P. 1861. Essai sur la faune entomologique de la Nouvelle-Calédonie (Balade) et des Iles des Pins, Art, Lifu, etc. Hémiptères. Annales de la Société Entomologique de France 1: 59–74. Motschulsky V. 1863. Essais d’un catalogue des insectes de l’ile Ceylan (Suite). Bulletin de la Société Impériale des Naturalistes de Moscou 36: 1–533. Mulsant E, Rey C. 1852. Description de quelques Hémiptères Hétéroptères nouveaux on peu connus. Annales de la Société Linnéenne de Lyon 1850/1852: 76–141. Namyatova A, Konstantinov F. 2009. Revision of the genus Orthocephalus Fieber, 1858 (Heteroptera: Miridae: Orthotylinae). Zootaxa 2316: 1–118. Nonnaizab SK, Bao A-P. 2006. A new record genus and a new species of Orthotylinae from China (Hemiptera, Heteroptera, Miridae). Acta Zootaxonomica Sinica 31: 585–587. Oshanin B. 1910. Verzeichnis der Palaearktischen Hemipteren. Kaiserlichen Akademie der Wissenschaften, St. Petersburg 3: 1–218.

Oshanin B. 1912. Katalog der paläarktischen Hemipteren. Berlin: Friedländer & Sohn. Panzer GWF. 1798. Fauna Insectorum Germaniae initia oder Deutschlands Insecten gesammelt und herausgegeben. Heft 49–60. Nürnberg: Felssecker. Panzer GWF. 1804. Faunae insectorum Germanicae initia oder Deutschlands Insecten gesammelt und herausgegeben. Heft 86–96. Nürnberg: Felssecker. Pericart J. 1965. Contribution à la faunistique de la Corse: Héteroptères Miridae et Anthocoridae (Hem.). Bulletin Mensuel de la Société Linnéenne de Lyon 34: 377– 384. Pluot-Sigwalt D, Matocq A. 2006. On some particular sclerotized structures associated with the vulvar area and the vestibulum in the Orthotylinae and Phylinae (Heteroptera, Miridae). Denisia 19: 557–570. Poppius B. 1910. 12. Hemiptera 4. Miridae, Anthocoridae, Termatophylidae, Microphysidae, und Nabidae. In: Sjöstedt Y, ed. Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Afrikas, vol. 2. Stokholm: Palmqvist, 8–14, 25–60. Poppius B. 1911. Beiträge zur Miriden-Fauna Australiens. Öfversigt af Finska Vetenskaps-Societetens Förhandlingar 53A: 1–6. Poppius B. 1914. Die Miriden der Äthiopischen Region II – Macrolophinae, Heterotominae, Phylinae. Acta Societatis Scientarum Fennicae 44: 1–138. Provancher L. 1887. Petite Faune Entomologique du Canada et Particulièrement de la Province de Québec. Vol. 3, Cinquième ordre. Les Hémiptères. Québec: C. Darveau. Puton A. 1874. Notes pour servir à l’étude des Hémiptères. 2ème Partie. Annales de la Société Entomologique de France 5: 213–314. Puton A. 1886. Catalogue des Hémiptères (Hétéroptères, Cicadines et Psyllides) de la faune Paléarctique. Caen: Henri Delesques. Puton A. 1887. Hémiptères nouveaux de l’Algerie. Revue d’Entomologie 6: 298–311. Puton A. 1895. Hémiptères nouveaux. Revue d’Entomologie 14: 83–91. Puton A. 1898. Quatre Hémiptères nouveaux. Revue d’Entomologie 17: 273–274. Putshkov VG. 1961. A new genus and two new species of the tribe Halticini Kirk. (Miridae, Heteroptera) from East Ukraine. Zoologicheskii Zhurnal 40: 25–30. Rambur JP. 1839. Faune entomologique de l’Andalousie. Paris: Arthur Bertrand, 97–176. Reuter OM. 1875a. Revisio critica Capsinarum, praecipue Scandinaviae et Fenniae. Försök till de Europäiska Capsinernas naturelinga uppställning jämte kritisk öfversigt af de Skandinavisk-finska arterna. Akademisk Afhandling Helsingfors 1: 1–101. Reuter OM. 1875b. Genera Cimicidarum Europae. Bihang till Kongliga Svenska Vetenskapsakademiens Forhandlingar 3: 1–66. Reuter OM. 1875c. Hemiptera gymnocerata scandinaviae et fenniae disposuit et descripsit. Pars i cimicidae (capsina).


THE HALTICINI OF THE WORLD Helsingfors: Ex officina typographica societatis litterariae fennicae. Reuter OM. 1879. Diagnoses Hemipterorum novorum. Öfversikt af Finska Vetenskaps-Societetens Förhandlingar 21: 30–41. Reuter OM. 1880. Diagnoses Hemipterorum novorum. II. Öfversigt af Finska Vetenskapssocietetens Förhandlingar 22: 9–24. Reuter OM. 1881. Acanthosomina et Urolabidina. Nova et minus cognita. Berliner Entomologische Zeitschrift 25: 67–86. Reuter OM. 1882. Labops putoni, n. sp. Capsidarum e Gallia descripsit. Revue d’Entomologie 1: 277–278. Reuter OM. 1883. Hemiptera Gymnocerata Europae. Hémiptères Gymnocèrates d’Europe, du bassin de la Méditerranée et de l’Asie Russe. III. Acta Societatis Scientiarum Fennicae 13: 313–496. Reuter OM. 1885. Species Capsidarum quas legit expeditio danica Galateae descripsit. Entomologisk Tidskrift 5: 195– 200. Reuter OM. 1888. Revisio synonymica Heteropterorum palaearcticorum quae descripserunt auctores vetustiores (Linné 1758-Latreille 1806) beschrieben Palaearktischen Heteropteren. Acta Societatis Scientiarum Fennicae 15: 241–315, 443–812. Reuter OM. 1890. Adnotationes Hemipterlogicae. Revue d’Entomologie 9: 248–254. Corrigenda 10: 27 (1891). Reuter OM. 1891. Hemiptera Gymnocerata Europae. Hémiptères Gymnocèrates d’Europe du bassin du Méditerranée et de l’Asie Russe. Tome IV. Acta Societatis Scientiarum Fennicae 23: 1–179. Reuter OM. 1894. Ad cognitionem Capsidarum. II Capsidae palaearcticae. Revue Française d’Entomologie 13: 128–152. Reuter OM. 1895. Ad cognitionem Capsidarum. III. Capsidae ex Africa boreali. Revue d’Entomologie, Caen 14: 131–142. Reuter OM. 1899. Capsidae novae mediterranae. I. Öfversigt af Finska Vetenskaps-Societetens Förhandlingar 42B: 131– 162. Reuter OM. 1901. Capsidae rossicae descriptae. Öfversigt af Finska Vetenskaps-Societetens Förhandlingar 45: 161–194. Reuter OM. 1903. Capsidae novae Rossicae. II. Öfversigt af Finska Vetenskaps-Societetens Förhandlingar 46: 1–17. Reuter OM. 1904. Ad cognitionem Capsidarum aethiopicarum. Öfversikt af Finska Vetenskaps-Societetens Förhandlingar 46: 1–8. Reuter OM. 1905a. Hemipterologische Spekulationen. I: Die Klassifikation der Capsiden. Festschrift fur Palmen No.1. Helsinfors. Reuter OM. 1905b. Descriptions of two new species of Hemiptera-Heteroptera from Spain. Transactions of the Entomological Society of London 1905: 52–54. Reuter OM. 1905c. Trois Capsides nouveaux pour la faune française (Hem). Bulletin de la Société Entomologique de France 1905: 8–9. Reuter OM. 1906. Capsidae in prov. Sz’tschwan Chinae a DD. G. Potanin et M. Beresowski collectae. Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg 10: 1–81.

649

Reuter OM. 1908. Capsidae novae palaearcticae. Annuaire du Musee Zoologique, St. Petersburg 12: 484–499 (1907). Reuter OM. 1909. Charakteristik und Entwicklungsgeschichte der Hemipterenfauna (Heteroptera, Auchenorrhyncha und Psyllidae) der palaearktischen Coniferen. Acta Societatis Scientiarum Fennicae 36: 1–129. Reuter OM. 1910. Neue Beiträge zur Phylogenie und Systematik der Miriden nebst einleitenden Bemerkungen über die Phylogenie der Heteropteren-Familien. Acta Societatis Scientiarum Fennicae 37: 1–167. Ribes J. 1984. Troballes noves o remarcables d’Hemipters per a Catalunya. Sessió Conjunta d’Entomologia ICHN-SCL 3: 105–115. Ribes J, Pagola-Carte S. 2007. Brachynema purpureomarginatum kerzhneri n. sp. from Iran (Hemiptera: Heteroptera: Pentatomidae), with a key to the ‘Rassenkreis’. Heteropterus Revista de Entomología 7: 19–24. Ribes J, Ribes E. 2001. A new species of Dimorphocoris Reuter, 1890 from Morocco (Heteroptera, Miridae). Bulletin de la Société Entomologique de France 106: 423– 425. Sahlberg J. 1875. Hemiptera Heteroptera nova fennica descripsit. Notiser ur Sällskapets pro Fauna et Flora Fennica Förhandlingar 14: 305–312. Sahlberg J. 1878. Bidrag till Nordvestra Sibiriens Insektfauna, Hemiptera-Heteroptera. Kungliga Svenska Vetenskapsakademiens Handlingar 16: 1–39. Saunders E. 1875. Synopsis of British HemipteraHeteroptera. Part I. Transactions of the Royal Entomological Society of London 1875: 117–129. Saunders E. 1892. The Hemiptera-Heteroptera of the British Islands. A descriptive account of the families, genera and species indigenous to Great Britain and Ireland, with notes as to localities, habitats, etc. London: L. Reeve & Co. Saunders E. 1894. A list of the Hemiptera-Heteroptera collected by Mr. Champion in Corsica in June, 1893, with a description of one new species. Transactions of the Royal Entomological Society of London 1894: 243–247. Say T. 1832. Descriptions of new species of Heteropterous Hemiptera of North America. New Harmony, Indiana. Schaffner JC, Schwartz MD. 2008. Revision of the Mexican genera Ficinus Distant and Jornandes Distant, with the description of 21 new species (Heteroptera, Miridae, Orthotylinae, Orthotylini). Bulletin of the American Museum of Natural History 309: 1–87. Schmidt K. 1939. Ein neuer Strongylocoris Blanch. aus Portugal. (Hem. Het. Laboparia). Memorias e Estudos do Museu Zoológico da Universidade de Coimbra 113: 1. Schuh RT. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with a phylogenetic analysis of the ant-mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1–332. Schuh RT. 1975. The structure, distribution and taxonomic importance of trichobothria in the Miridae (Hemiptera). American Museum Novitates 2585: 1–22. Schuh RT. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of relationships within the family. American Museum Novitates 2601: 1–39.


650


Schuh RT. 1995. Plant bugs of the world (Insecta: Heteroptera: Miridae). Systematic catalog, distributions, host list, and bibliography. New York: The New York Entomological Society. Schuh RT, Lattin JD. 1980. Myrmecophyes oregonensis, a new species of Halticini (Hemiptera, Miridae) from the western United States. American Museum Novitates, 2697: 1–11. Schuh RT, Slater JA. 1995. True bugs of the world (Hemiptera: Heteroptera): classification and natural history. Ithaca, NY: Cornell University Press. Schuh RT, Weirauch C, Wheeler WC. 2009. Phylogenetic relationships within the Cimicomorpha (Hemiptera: Heteroptera): a total-evidence analysis. Systematic Entomology 34: 15–48. Schwartz MD, Schuh RT, Tatarnic N. 2008. A new genus and species of Halticini from South Africa (Hemiptera: Miridae). African Entomology 16: 23–32. Scott J. 1870. Neue europäische Hemiptera. Stettiner Entomologische Zeitung 31: 98–102. Scott J. 1874. Description of a genus and species of Hemiptera-Heteroptera new to Europe. Entomologist’s Monthly Magazine 10: 272–273. Seidenstücker G. 1956. Eine neue palaearktische Miriden mit Membrane-Pubeszenz (Hemiptera-Heteroptera). Review of the Faculty of Science, University of Istanbul (Series B) 21: 65–70. Seidenstücker G. 1959. Heteroptera aus Anatolien II. Review of the Faculty of Science, University of Istanbul (Series B) 23: 119–129. Seidenstücker G. 1962. Über einige Miriden aus Kleinasien mit Beschreibung von zwei neuen Halticinen (Heteroptera). Reichenbachia 1: 129–143. Seidenstücker G. 1971. Zwei neue Orthotylinae (Heteroptera, Miridae). Nachrichtenblatt der Bayerischen Entomologen 20: 36–40. Signoret V. 1862. Coridromius n. n. f. Ocypus Montrouzier. Bulletin de la Société Entomologique de France 4: 5. Simov N. 2005. A new Scirtetellus species from the Karakoram Mts., Pakistan (Heteroptera: Miridae). Zoosystematica Rossica 14: 213–214. Slater JAS. 1950. An Investigation of the female genitalia as taxonomic characters in the Miridae (Heteroptera). Iowa State College Journal of Science 25: 1–81. Slater JAS. 1954. Notes on the genus Labops, Burnmeister in North America, with the descriptions of three new species (Hemiptera: Miridae). Bulletin of the Brooklyn Entomological Society 49: 1–15. Southwood TRE. 1953. The morphology and taxonomy of the genus Orthotylus Fieber (Hemiptera, Miridae), with special reference to British species. Transactions of the Royal Society of London 104: 415–499. Southwood TRE. 1962. Migration of terrestrial arthropods in relation to habitat. Biological Reviews 37: 171–214. Southwood TRE, Leston D. 1959. Land and water bugs of the British Isles. London: Frederick Warne and Co. Spinola MM. 1840. Essai sur les insectes Hémiptères, Rhyngotes ou Hétéroptères. Paris: Baillière.

Spinola MM. 1850. Tavola sinottica dei generi spettanti alla classe degli Insetti artroidignati, Hemiptera, Linn. Latr. – Rhyngota, Fab. – Rhynchota Burm. Memoria del Socio Attuale signor Marchese Massimiliano Spinola Modena, Dal tipi delle R.D. Camera. Societa Italiana delle Scienze 25: 43–100. Stål C. 1858. Beitrag zur Hemipteren-Fauna Sibiriens und des Russischen Nord-Amerika. Stettiner Entomologische Zeitung 19: 175–198. Stichel W. 1933. Illustrierte Bestimmungstabellen der Deutschen Wanzen (Hemiptera-Heteroptera). Fasc. 8, 9. BerlinHermsdorf: W. Stichel. Sweet MH. 1964. The biology and ecology of the Rhyparochrominae of New England (Heteroptera: Lygaeidae) Part I. Entomologia Americana 43: 1–124. Tamanini L. 1955. Descrizione di un nuovo miride Plagiotylus zorzii n. sp. (Hemiptera Heteroptera, Miridae). Memorie del Museo Civico di Storia Naturale di Verona 5: 31–38. Tamanini L. 1960. Descrizione di un nuovo Plagiotylus per la Sicilia. Con alcune osservazioni sulla distribuzione delle altre specie. (Hemiptera Heteroptera, Miridae). Memorie del Museo Civico di Storia Naturale di Verona 8: 79–88. Tamanini L. 1971. Un emittero nuovo delle Alpi italiane (Hemiptera Heteroptera, Miridae). Studi Trentini di Scienze Naturali (Ser. B) 48: 500–511. Tamanini L. 1972. Descrizione di due nuovi Dimorphocoris montani utilizzando anche i caratteri delle armature endofalliche (Hemiptera Heteroptera Miridae). Atti Della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 113: 117–132. Tamanini L. 1975. Un nuovo Dimorphocoris deele Alpi Cozie con ulteriori notizie sul Dimorphocoris ruffoi Tam., 1971. Bollettino del Museo Civico di Storia Naturale di Verona 2: 323–330. Tamanini L. 1976. Dimorphocoris puigmalis n.sp., dei pirenei orientali (Hemiptera, Heteroptera, Miridae). Bollettino della Società Entomologica Italiana 108: 147–151. Tamanini L. 1981. Dimorphocoris servadeii n. sp. degli appennin: (Heteroptera Miridae). Memorie della Societa Entomologica Italiana 60: 335–341. Tamanini L. 1982. Dimorphocoris servadeii n. sp., degli Appennini (Heteroptera, Miridae). Memorie della Società Entomologica Italiana 60: 335–341. Tatarnic NJ. 2009. Dampierella and Goodeniaphila: two new genera and three new species of Halticini from Australia, with a species key to the Halticini of Australia (Hemiptera: Heteroptera: Miridae: Orthotylinae). Zootaxa 2105: 43– 60. Tatarnic NJ, Cassis G. 2008. Revision of the plant bug genus Coridromius Signoret (Insecta: Heteroptera: Miridae). Bulletin of the American Museum of Natural History 315: 1–95. Tatarnic NJ, Cassis G. 2010. Sexual coevolution in the traumatically inseminating plant bug genus Coridromius. Journal of Evolutionary Biology 23: 1321–1326. Tatarnic NJ, Cassis G, Hochuli DF. 2006. Traumatic insemination in the plant bug genus Coridromius Signoret (Heteroptera: Miridae). Biology Letters 2: 58–61.


THE HALTICINI OF THE WORLD Thompson CG. 1871. Opuscula entomologica. Hemiptera, Fasc IV. Lund. Uhler PR. 1872. Notices on the Hemiptera of the western territories of the United States, chiefly from the surveys of Dr. F. V. Hayden. In: Heydon FV, ed. Preliminary report of the United States Geological survey of Montana and portions of adjacent territories, being a fifth annual report of progress. Washington, DC: Government Printing Office, 392–423. Uhler PR. 1904. List of Hemiptera-Heteroptera of Las Vegas, Hot Springs, New Mexico, collected by Messrs. E. A. Schwarz and Herbert S. Barber. Proceedings of the United States National Museum 27: 349–364. Usinger RL. 1946. Hemiptera Heteroptera of Guam. In: Swezey OH, ed. Insects of Guam, Vol II. Bulletin of the Bishop Museum No. 189. Honolulu: Bishop Museum Press, 11–103. Van Duzee EP. 1914. Nomenclatorial and critical notes on Hemiptera. Canadian Entomologist 46: 377– 389. Van Duzee EP. 1916. Check list of the Hemiptera (excepting the Aphididae, Aleurodidae and Coccidae) of America, North of Mexico. New York: New York Entomological Society. Van Duzee EP. 1917. Catalogue of the Hemiptera of America north of Mexico, excepting the Aphididae, Coccidae and Aleurodidae. University of California College of Agriculture, Agricultural Experiment Station, Technical Bulletin, Entomology 2: 1–902. Van Duzee EP. 1921. Our North American species of Strongylocoris (Hemiptera). Proceedings of the California Academy of Sciences 11: 135–136. Villiers A. 1952. Initiations Africaines. IX. Hémiptères de l’Afrique Noire (Punaises et cigales). Institute Francaise d’Afrique Noire, Dakar 9: 1–256. Vinokurov NN. 1979. Identification manual of the fauna of the USSR. No. 123. Heteroptera of the Yakutia. Leningrad: Zoological Institute of the USSR. Wagner E. 1940. Zur Systematik von Lygus pratensis L. (Hemiptera, Heteroptera, Miridae). Ebenda 28: 1–6. Wagner E. 1951. Halticus major n. sp., eine neue Miridenart. Nachrichten des Naturwissenschaftlichen Museums der Stadt Aschaffenburg 33: 69–75. Wagner E. 1952. Blindwanzen oder Miriden. Die Tierwelt Deutschlands 41: 1–218. Wagner E. 1954. Eine neue Miriden-Art aus Sud-Frankreich (Hemipt. Heteropt.). Revue Française d’Entomologie 21: 219–223. Wagner E. 1955. Bemerkungen zum System der Miridae (Hem. Het.). Deutsche Entomologische Zeitschrift, N. F. 2: 230–242. Wagner E. 1956. La validité de l’espèce Strongylocoris oberthuri Reuter. Vie et Milieu 7: 101–103. Wagner E. 1957. Zwei neue Dimorphocoris-Arten aus Frankreich (Hem. Het. Miridae). Deutsche Entomologische Zeitschrift, N.F. 4: 223–227. Wagner E. 1960. Was ist Dasyscytus sordidus Fieber 1864? (Hem. Het. Miridae). Mitteilungen der Deutsche Entomologischen Gesellschaft 19: 98–101.

651

Wagner E. 1961. Unterordnung: Ungleichflugler, Wanzen, Heteroptera (Hemiptera). Die Tierwelt Mitteleuropas 4: 1–173. Wagner E. 1965a. Uber die Gattung Dimorphocoris Reuter, 1891 (Hemiptera, Heteroptera, Miridae). Reichenbachia 5: 135–156. Wagner E. 1965b. Die Gattung Dimorphocoris Reuter, 1891, II (Hem. Het. Miridae). Reichenbachia 6: 33–66. Wagner E. 1966. Uber Dimorphocoris robustus E. Wagner, 1957 (Hemiptera, Heteroptera, Miridae). Reichenbachia 8: 71–74. Wagner E. 1968. Über eine neue und eine schon bekannte Miridengattung (Heteroptera, Miridae). Reichenbachia 10: 221–226. Wagner E. 1969. Uber Dimorphocoris saulii E. Wagner, 1965 (Hemiptera, Heteroptera, Miridae). Bollettino della Società Entomologica Italiana 99–101: 77–80. Wagner E. 1973. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Entomologishe Abhandlungen 39: 1– 423. Wagner E. 1975. Vier neue Miriden (Insecta, Hemiptera, Heteroptera) aus dem Mittelmeergebiet. Bulletin Zoölogisch Museum Universiteit van Amsterdam 4: 141–148. Wagner E. 1976. Noch zwei Myrmecophyes-Arten aus dem Mittelmeerraum (Hemiptera, Heteroptera, Miridae). Reichenbachia 16: 157–162. Wagner E, Weber HH. 1964. Hétéroptères Miridae. Faune de France 67: 1–592. Walker F. 1873. Catalogue of specimens of Hemiptera Heteroptera in the collection of the British Museum. Part VI. London: The British Museum. Westwood JO. 1840. An Introduction to the Modern Classification of Insects, founded on the natural habits and corresponding organisation of the different families. Synopsis of the genera of British insects, vol. II. London: Longman. Zetterstedt JW. 1838. Insecta Lapponica descripta. Leipzig: L. Voss. Zimmerman EC. 1948. Insects of Hawaii. Heteroptera, vol. III. Honolulu: University of Hawaii Press. Zou HG. 1985. Three new species of Halticus Hahn from China (Hemiptera: Miridae). Acta Zootaxonomica Sinica 10: 304–308.

APPENDIX 1 LIST

OF CHARACTERS AND CHARACTER STATES

(Figures cited refer to examples of structures described.) 1. Coloration: (0) variably coloured, not mostly black; (1) mostly black or dark brown. 2. Texture – head and thorax: (0) without patches of scale-like or honeycombed texturing; (1) with symmetrical patches of scale-like honeycombed texturing laterally on thorax and dorsally on head (Figs 15A, C, D, 22E).


652


3. Punctures – body and hemelytra: (0) impunctate, or with very few punctures; (1) nearly entirely punctate (Figs 5A–C, E, 43A–C, E). 4. Rugosity – pronotum: (0) not rugose (Figs 5A, 20A, 22A, 50B); (1) wholly or partly rugose (Figs 7A, 11A, 26A, B, 28A, 39A, 52B). 5. White scale-like or feather-like setae: (0) absent; (1) present on head, thorax, hemelytra or legs (Fig. 3: Anapomella arnoldi, Dampierella schwartzi; Fig. 17A–D). 6. Erect or semi-erect bristle-like setae on dorsum: (0) absent, or with only a few erect or semi-erect bristle-like setae; (1) dorsum clothed in erect or semi-erect dark, bristle-like setae (Fig. 41A: coloration not visible in scanning electron micrograph); (2) dorsum clothed in erect or semi-erect pale, bristle-like setae. 7. Texture – frons: (0) without radiating folds; (1) with radiating folds diverging anteriorly (Figs 26B–C, 39B). 8. Vertex sinuation (viewed anteriorly): (0) dorsal margin of vertex appearing flat or convex, but never concave (Figs 5C, 11B, 20B, 24B, 26C, 37B, 43B, 45B, 57B); (1) concave (Figs 33B, 35A, B). 9. Vertex – posterior margin: (0) rounded (Figs 33A, 39A, 54A); (1) sharply edged (Fig. 45A, C);. 10. Head shape as viewed from above: (0) transverse (much wider than long) (Figs 5A, 57B); (1) elongate (subequal, equal, or longer than wide) (Fig. 15C). 11. Paired, textured spots on vertex, superficially similar to ocelli: (0) absent; (1) present (Fig. 52B). 12. Conical projection of frons: (0) not conically projecting; (1) conically projecting (Fig. 13A, B). 13. Head height below eyes: (0) significantly less than height of eye; (1) less than or equal to height of eye (Figs 13B, 43D); (2) greater than height of eye but less than twice height of eye (Figs 37C, 47A); (3) greater than twice as high as the eye (Figs 33B, C, 39C). 14. Labial length: (0) reaching procoxa; (1) reaching mesocoxa; (2) reaching metacoxa; (3) reaching abdomen (surpassing metacoxa). 15. Angle of clypeus (viewed laterally): (0) appearing vertical or nearly so (Figs 24D, 33C); (1) distinctly recurved and angled posteriorly (Fig. 39C). 16. Stalked eyes: (0) eyes sessile; (1) eyes stalked (Fig. 33A–C). 17. Antennal fossa: (0) in line with medial portion of eye (Fig. 5B); (1) in line with lower margin of eye (Figs 17B-C, 28B, C); (2) below eye (Fig. 47A). 18. AI spines: (0) smooth or with minute spines only (Fig. 37A–C); (1) with a few long spines (Fig. 47A); (2) with multiple long spines (Figs 7A–C, E, 26A-E).

19. AI length: (0) less than height of eye (Figs 17B, C, 28B); (1) equal or slightly greater than height of eye (Fig. 37B, C); (2) much longer than height of eye (Figs 13A, 33B, C). 20. AIV length: (0) shorter or equal to AIII; (1) longer than AIII. 21. Total antennal length: (0) subequal to body length; (1) longer than body. 22. Metathoracic spiracle shape: (0) narrow and slitlike; (1) teardrop-shaped (Fig. 39D). 23. Metathoracic scent gland: (0) with MTG opening, peritreme and evaporative area all well developed (Figs 11D, E, 39D, E); (1) some or all components of MTG efferent system reduced or obsolete. 24. Metathoracic scent gland – size of evaporative area: (0) evaporative area minute or absent (Figs 47C, D, 50C, D, 57C, D); (1) small, occupying no greater than half metepimeron (Fig. 5E, F); (2) large, extending nearly to dorsum (Figs 11D, E, 26F). 25. Metathoracic scent gland – orientation of peritreme: (0) vertical (Fig. 26F); (1) angled somewhat caudally (Fig. 33C, D); (3) angled back and resting along caudal margin of tergite (Figs 5E, F, 43E, F). 26. Metathoracic scent gland – peritreme shape: (0) long and narrow or teardrop-shaped (Figs 5E, F, 33D, 39D, 43E, F); (1) broadly elongate-oval or round (Figs 37D, E, 52E). 27. Angle of proepisternum (viewed laterally): (0) surface of proepisternum immediately adjacent to anterior thoracic margin laterally orientated (Fig. 33C); (1) surface of proepisternum anteriorly orientated and appressed to posterior of head (Figs 30D, 37C). 28. Pronotal shape (macropterous): (0) trapezoidal (Figs 5A, 17A, 24A); (1) campanulate (Figs 22A, 33A). 29. Pronotal collar: (0) absent or vestigial (Fig. 7A); (1) present in at least one sex (Fig. 20A). 30. Posterior margin of pronotum: (0) straight (Fig. 39A); (1) concave (Fig. 20A); (2) convex (Figs 37A, 57A). 31. Mesoscutum (macropterous): (0) not visible from above; (1) visible from above. 32. Mesoscutum (brachypterous): (0) not visible from above; (1) visible from above. 33. Hemelytral membrane cells: (0) two cells; (1) single cell (Figs 17A, 28A); (2) no cells. 34. Hemelytral polymorphism – male: (0) always or nearly always macropterous, with all components present; (1) polymorphic, with both macropterous and reduced-wing morphs; (2) always with reduced wings. 35. Hemelytra - medial fracture (reduced wing): (0) absent; (1) short (subequal to length of claval



36. 37.

38.

39. 40. 41. 42.

43. 44. 45. 46.

47.

48.

49. 50.

51.

52.

suture); (2) elongate (greater than length of claval suture). Hemelytra- cuneus (reduced-wing): (0) absent; (1) present. Hemelytra – degree of female brachyptery: (0) macropterous – all components present; (1) all components present but membrane reduced to a thin strip; (2) reduced wing (brachypterous, coleopterous, staphylinoidous) (Fig. 7A); (3) micropterous (wing pads) (Fig. 39A). Hemelytral length – (reduced-wing female): (0) exposing fewer than two tergites; (1) exposing two to three tergites; (2) exposing more than three tergites. Hemelytra – membrane setae: (0) naked; (1) covered with dense setae. Hemelytra – lateral corial margin: (0) straight or convex; (1) sinuate. Hind femur thickness – female: (0) thin; (1) incrassate. Pretarsus – fleshy pulvilli: (0) underside of claws without fleshy pulvilli (Figs 39F, 45E, 47F); (1) underside of claws with fleshy pulvilli (Figs 7F, 11F, 17H, 24F). Foretibiae – shape: (0) not apically swollen; (1) apically swollen and laterally compressed. Metatibia – spines: (0) without spines; (1) spinose. Metafemora – spines: (0) absent; (1) present. Metafemora – apex: (0) medially and laterally without rounded tumescences at apex; (1) medially and laterally with round, smooth, and slightly swollen tumescence at apex. Pygophore – posteroventral margin: (0) symmetrical or only weakly asymmetrical (Figs 37G, H, 39H, 45G, 47H); (1) asymmetrical, with deep concavity below left paramere (Figs 7E, 33F–H); (2) asymmetrical and complex [significantly rotated, armed with spicules, etc. (outgroup only)]. Left and right paramere – base: (0) neither paramere with elongate base; (1) both parameres with elongate and narrow base (Figs 9B, C, 12B, C); (2) base or right paramere elongate, that of left paramere much shorter (Figs 18A, B, 29A, B). Right paramere – apex: (0) not clubbed and cupped; (1) clubbed and cupped. Right paramere - length: (0) much shorter than left; (1) slightly shorter to slightly longer than left; (2) much longer than left. Extension of right paramere: (0) not extending or only slightly extending from pygophore; (1) extending considerably from pygophore (Figs 30H, 33G, H, 47G, H). Left paramere - shape: (0) more or less L-shaped; (1) trifurcate (Fig. 25B); (2) twisted and elongate (Fig. 31D).

653

53. Left paramere - sensory lobe: (0) without discernible sensory lobe (Figs 31D, 40B, 48C); (1) sensory lobe present but poorly developed (Figs 23B, 46C); (2) sensory lobe large and conspicuous (Figs 9B, 12B, 27A). 54. Left paramere - apophysis: (0) without recurved apical hook or knob (Fig. 6B); (1) with small downturned hook or knob (Figs 9B, 12B); (2) with bifurcate, hooked tip (Figs 14B, 19B). 55. Coupling of aedeagus and left paramere: (0) not coupled to form a piercing intromittent organ; (1) coupled to form a piercing intromittent organ. 56. Phallotheca – apex: (0) without laterally projecting apical flange; (1) with prominent laterally projecting apical flange (Fig. 36A). 57. Phallotheca – apical constriction: (0) not laterally constricted; (1) abruptly laterally constricted beyond base (Fig. 44A). 58. Ductus seminis – length and width: (0) relatively thick and stout, sometimes short (Figs 18C, 31B, 44A, B, 46A); (1) slender, long, and sinuous, frequently convoluted (Figs 9A, 14A, 16A, B, 21A, 49A). 59. Ductus seminis – ribbing: (0) present (Fig. 18C); (1) absent (Fig. 44B). 60. Secondary gonopore – apical extension: (0) does not extend as a sclerotized tube running to the apex of the phallotheca; (1) extends as a sclerotized tube to the apex of the phallotheca (Figs 6A, 44A, B). 61. Secondary gonopore – apical sclerotization: (0) with a short or long sclerotized section beginning subapical to opening (Figs 8A, 55A); (1) mostly unsclerotized, if sclerotized then only along rim of secondary gonopore (Figs 16B, 21A). 62. Ductus seminis – apical sclerotization: (0) apical sclerotized region of the ductus seminis continuous, unbroken by clear transverse band (Fig. 40A); (1) apical sclerotization of the ductus seminis broken by clear transverse band prior to the secondary gonopore (Fig. 48B). 63. Ductus seminis – length of apical sclerotization: (0) apical sclerotized region short, barely extending below the secondary gonopore (Figs 46A, 53A, 55A); (1) apical sclerotized region of intermediate length (Figs 18C, 34A, 40A); (2) apical sclerotized region elongate (Figs 8A, 9A, 27C). 64. Secondary gonopore – width (when viewed from above): (0) narrow; (1) broad (Fig. 44A). 65. Secondary gonopore – dorsoventral compression: (0) not dorsoventrally compressed, more or less cylindrical; (1) dorsoventrally compressed (Fig. 46B). 66. Secondary gonopore – shape: (0) forming a narrow scoop (Figs 18C, 31B, C); (1) not forming a narrow scoop.


654


67. Secondary gonopore – ventral wall: (0) ventral wall obsolete, not distinctly recurved; (1) ventral wall distinctly recurved, forming a prised operculum (Figs 9A, 51A, 55A, 58C). 68. Secondary gonopore – subapical shape: (0) without distinct subapical bowl-shaped constriction; (1) with distinct sub-apical bowl-shaped constriction and funnel-shaped opening (Fig. 46A, B). 69. Secondary gonopore – shape of opening: (0) opening of secondary gonopore oval; (1) opening of secondary gonopore slit-like. 70. Secondary gonopore – texture: (0) smooth; (1) with scale-like texture (Fig. 46B). 71. Endosoma: (0) reduced or absent (Fig. 31B, C); (1) forming a well-developed membranous sac (Figs 34A, 36B). 72. Endosoma – spicules: (0) absent; (1) present. 73. Endosoma – spicule location: (0) attached basal to secondary gonopore; (1) distal to secondary gonopore. 74. Endosoma – spicule form: (0) without paired sclerites attached to anterior of secondary gonopore; (1) with paired paddle-like sclerites extending from anterior of secondary gonopore (Figs 16A, B, 23A, 53A). 75. Endosoma – spicule form: (0) when present, simple (Figs 9A, 12A); (1) spicules, when present complex and branching (Figs 29C, D, 38A). 76. Endosoma – spicule form: (0) without tight bunch of spinose spicules; (1) with tight bunch of spinose spicules (Figs 34A, 36B). 77. Endosoma – fields of spines: (0) without fields of spines; (1) apically with sclerotized fields of spines (Figs 19A, 51A); (2) apically with one or two discrete, elongate sclerites covered with fields of spines (Figs 34A, 40A). 78. Female - sclerotized rings: (0) present; (1) absent. 79. Female - sclerotized rings: (0) widely separated; (1) medially contiguous or subcontiguous. 80. Female - sclerotized rings: (0) no larger than diameter of posterior wall; (1) larger than diameter of posterior wall.

81. Ventral labiate plate – region bordering rami: (0) not forming a sclerotized rim; (1) forming a moderately to strongly sclerotized rim (Fig. 9E). 82. Inter-ramal bridge: (0) absent; (1) present (Figs 10A, 34G, 36E, F). 83. Dorsal labiate plate – medial sclerites: (0) absent; (1) with paired, sickle-shaped sclerites adjacent to sclerotized rings. 84. Rami – medial projections: (0) absent; (1) present (Figs 34G, 35A). 85. Posterior wall – sclerotization: (0) wholly or partly sclerotized; (1) entirely membranous (Figs 6F, 14E, 44H). 86. Posterior wall – texture: (0) with fields of spines (Figs 34H, 44H); (1) without fields of spines (Fig. 6F). 87. Posterior wall – structure: (0) without bilateral pillow-like swellings; (1) with bilateral pillow-like swellings (Figs 6F, 25E, 44H). 88. Posterior wall – inter-ramal lobes: (0) absent; (1) with prominent, paired, projecting spinose or serrate lobes (Figs 8E, 9F, 10B, 34H, 35B, 36G, 55D, 55C). 89. Posterior wall – base of inter-ramal lobes: (0) paired, hollow inter-ramal lobes narrow and strongly constricted basally (outgroup only); (1) inter-ramal lobes not narrow and basally constricted. 90. Posterior wall – division: (0) distinctly tripartite, with discrete lateral plates above a smaller medial plate (outgroup only); (1) occasionally bilaterally modified, but without discrete lateral and basal plates. 91. Opening to vestibulum- symmetry of first gonapophyses: (0) asymmetrical; (1) symmetrical (Figs 6E, 10C, 25G). 92. Opening to vestibulum – degree of sclerotization: (0) entirely membranous (Fig. 14D); (1) with at least some sclerotization (Figs 6E, 10C).


655


APPENDIX 2 Halticini morphological data matrix.

Orthotylus sp. Ceratocapsus sp. Nichomachus minutus Aopolonema princeps Lattinova jacki Acratheus ocellaris Anapus americanus Anapus kirshbaumi Anapus rugicollis Anapus dorsalis Barbarosia punctulata Chorosomella jakowleffi Coridromius variegatus Dasyscytus sordidus Dicyphopsis sp. Compositocoris senecionus Dampierella schwartzi Dimorphocoris gracilis Dimorphocoris fuscus Ectmetopterus micantulus Euryopicoris nitidus Goodeniaphila cassisi Halticus apterus Halticus intermedius Halticus luteicollis Labops burnmeisteri Labops utahensis Labops sahlbergii Labops tumidiferons Microtechnites altigena Microtechnites bractatus Myrmecophyes alboornatus Myrmecophyes oregonensis Namaquacapsus melanostethoides Nanniella palustris Orthocephalus arnoldii Orthocephalus brevis Pachytomella passerini Piezocranum simulans Plagiotylus maculatus Schoenocoris flavomarginatus Scirtetellus seminitens Scirtetellus brevipennis Strongylocoris leucocephalus

Orthotylus sp. Ceratocapsus sp. Nichomachus minutus Aopolonema princeps Lattinova jacki Acratheus ocellaris Anapus americanus Anapus kirshbaumi Anapus rugicollis Anapus dorsalis Barbarosia punctulata Chorosomella jakowleffi Coridromius variegatus Dasyscytus sordidus Dicyphopsis sp. Compositocoris senecionus

1

2

3

4

5

6

7

8

9

1 0

1 1

1 2

1 3

1 4

1 5

1 6

1 7

1 8

1 9

2 0

0 0 0 1 0 1 1 1 1 1 1 0 0 1 0 0 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

0 0 1 0 0 0 1 1 1 1 1 0 0 1 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 0 1 1 1 0

0 0 0 ? 0 0 1 1 1 0 0 0 0 1 0 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 1 1 1 1 0

0 2 0 0 0 0 0 0 0 0 0 0 0 2 0 1 0 1 1 0 0 0 0 0 0 2 2 2 2 0 0 0 0 1 0 1 1 1 1 1 1 1 1 0

0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 1 1 0

0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0

0 1 1 1 0 1 0 0 0 0 0 0 1 1 1 0 1 0 0 1 0 1 1 1 1 0 0 0 0 1 1 0 0 1 1 1 1 1 1 1 0 0 0 1

1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 1 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 1 0 0 1 1 0 1 0 1 1 1 1 1 1 1 1

0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 0 0 0 0 2 3 3 3 3 3 1 1 2 1 1 1 2 3 2 3 1 2 2 2 3 3 3 3 2 2 3 3 2 1 2 2 2 2 2 2 3 3 2

2 2 2 ? 2 2 2 1 2 2 1 0 2 1 2 1 2 2 2 2 1 1 1 1 0 2 2 2 1 2 2 1 2 2 1 2 1 1 2 2 2 2 2 1

0 1 1 0 0 0 1 1 1 1 1 0 0 1 1 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 1 1 1 0 1 0 1 1 1 1 1 1 1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

2 2 1 1 1 1 3 3 3 3 3 1 1 3 1 2 2 3 3 1 3 2 1 1 1 3 3 3 3 1 1 3 3 2 1 3 3 3 3 3 3 3 3 3

0 1 0 0 1 0 2 2 2 2 1 0 0 1 1 2 1 2 2 1 2 0 1 1 1 2 2 2 2 0 0 1 1 0 0 0 1 1 0 1 2 2 2 1

1 0 0 ? 2 1 2 2 2 2 1 2 0 1 1 2 0 2 2 1 2 0 0 1 1 2 2 2 2 0 1 2 2 0 1 0 1 1 0 2 2 2 2 0

0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0

2 1

2 2

2 3

2 4

2 5

2 6

2 7

2 8

2 9

3 0

3 1

3 2

3 3

3 4

3 5

3 6

3 7

3 8

4 9

4 0

0 0 0 0 1 1 1 0 0 0 0 1 0 ? 1 1

1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1

0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0

1 1 1 1 1 1 1 1 1 1 2 0 1 1 1 1

1 2 1 1 1 3 1 1 1 1 1 3 1 3 1

1 0 0 0 0 0 1 1 1 1 1 0 0 0 0

1 1 1 1 1 0 0 0 0 0 1 0 1 1 0 0

0 0 0 0 1 0 0 0 0 0 0 0 1 1

0 1 1 1 0 1 0 0 0 0 0 0 1 ? 1 1

0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1

0 1 1 1 1 0 1 1 1 1 1 0 0 1

? 0 0 0 0 0 1 0 0

0 0 0 0 1 0 1 1 1 0 0 0 0

0 0 0 0 0 0 2 1 1 1 1 0 0 0 0 0

? 0 0 0 0 0 1 0 0

? 0 0 0 0 0 0 0 0

0 0 ? 0 0 0 3 2 2 2 2 3 0 2 0 3

? 2 2 1 2 1 2 2 2

0 0 0 0 0 0 0 0 0 0 0 0 1 0 0

0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0


656


APPENDIX 2

Dampierella schwartzi Dimorphocoris gracilis Dimorphocoris fuscus Ectmetopterus micantulus Euryopicoris nitidus Goodeniaphila cassisi Halticus apterus Halticus intermedius Halticus luteicollis Labops burnmeisteri Labops utahensis Labops sahlbergii Labops tumidiferons Microtechnites altigena Microtechnites bractatus Myrmecophyes alboornatus Myrmecophyes oregonensis Namaquacapsus melanostethoides Nanniella palustris Orthocephalus arnoldii Orthocephalus brevis Pachytomella passerini Piezocranum simulans Plagiotylus maculatus Schoenocoris flavomarginatus Scirtetellus seminitens Scirtetellus brevipennis Strongylocoris leucocephalus

Orthotylus sp. Ceratocapsus sp. Nichomachus minutus Aopolonema princeps Lattinova jacki Acratheus ocellaris Anapus americanus Anapus kirshbaumi Anapus rugicollis Anapus dorsalis Barbarosia punctulata Chorosomella jakowleffi Coridromius variegatus Dasyscytus sordidus Dicyphopsis sp. Compositocoris senecionus Dampierella schwartzi Dimorphocoris gracilis Dimorphocoris fuscus Ectmetopterus micantulus Euryopicoris nitidus Goodeniaphila cassisi Halticus apterus Halticus intermedius Halticus luteicollis Labops burnmeisteri Labops utahensis Labops sahlbergii Labops tumidiferons Microtechnites altigena Microtechnites bractatus Myrmecophyes alboornatus Myrmecophyes oregonensis Namaquacapsus melanostethoides

Continued

2 1

2 2

2 3

2 4

2 5

2 6

2 7

2 8

2 9

3 0

3 1

3 2

3 3

3 4

3 5

3 6

3 7

3 8

4 9

4 0

0 1 1 1 0 0 1 1 1 1 1 1 1 0 1 1 1 0 0 0 0 0 0 0 0 1 1 0

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 1 0 1 0 0 0 1

1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 0 1 0 1 1 1 0

1 2 2 3 1 1 1 1 1 2 2 2 2 1 1 1 1 ? 3 1 1 3 1 1 1 -

1 0 0 0 1 1 0 0 1 0 0 0 0 1 1 0 0 ? 0 1 1 0 1 0 0 -

1 0 0 0 0 1 1 1 1 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1 1 0 0 1

0 1 1 0 0 0 0 0 0 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 0

0 1 1 1 0 0 0 0 1 1 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 0 0

1 1 1 0 1 1 0 0 0 1 1 1 1 2 1 0 0 0 1 0 0 1 1 1 1 0 0 2

0 1 1 1 1 0 1 0 0 1 1 1 1 0 0 1 1 0 0 1 1 1 1 1 1 0

0 0 0 0 1 1 1 1 0 1 1 ? 0 0 0 0 1 0 0 0 -

1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 0 0 0 0 0

0 0 0 0 1 0 1 0 0 0 1 1 0 0 1 1 2 0 0 0 1 0 0 0 0 2 2 0

0 0 0 1 1 1 1 1 1 0 0 0 ? 0 1 0 0 1 0 0 0 -

0 0 0 1 1 1 1 1 1 0 0 0 ? 0 1 0 0 0 0 0 0 -

0 3 3 0 2 0 2 0 0 1 1 1 1 0 2 3 3 ? 0 2 2 2 2 2 2 3 3 0

2 2 1 1 0 0 0 0 1 2 2 ? 1 1 1 1 1 2 2 2 -

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

4 1

4 2

4 3

4 4

4 5

4 6

4 7

4 8

4 9

5 0

5 1

5 2

5 3

5 4

5 5

5 6

5 7

5 8

5 9

6 0

0 0 ? 0 0 0 1 1 1 1 1 0 1 0 1 0 1 0 0 1 1 1 1 1 1 0 0 0 0 1 1 1 1 0

0 1 0 1 0 0 0 1 1 1 1 1 1 0 0 1 1 0 0 1 1 1 1 1 1 0 0 0 0 1 0 0 0 1

0 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0

1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0

0 0 0 ? 1 0 1 1 1 1 1 0 0 0 0 1 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0

0 0 0 0 0 0 1 1 1 1 1 0 1 ? 0 0 0 0 0 0 1 0 1 1 1 0 0 0 0 1 1 1 1 0

2 1 1 2 2 0 1 1 1 1 0 0 1 0 0 1 0 0 0 1 0 0 1 1 1 1 1 1 1 0 0 0 0 ?

0 0 0 1 0 1 1 1 1 1 1 1 0 1 1 1 2 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1

0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 0 1 1 1 1 1 1 2 1 1 0 1 2 1 2 2 2 2 1 2 2 2 2 2 2 2 2 1 1 2 2 2

0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 0 0 0 1 ?

0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 2 2 0 0 0 0 0 0 0 0 0

1 2 2 0 2 1 0 2 0 2 2 1 0 2 1 1 2 1 1 1 2 2 0 0 0 2 2 2 2 2 2 0 0 2

0 2 1 1 0 0 1 1 1 1 1 2 0 2 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2

0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 1 1 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 0 0 0 0 1 1 1 1 0 0 1 1 1

0 0 1 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0


657


APPENDIX 2

Nanniella palustris Orthocephalus arnoldii Orthocephalus brevis Pachytomella passerini Piezocranum simulans Plagiotylus maculatus Schoenocoris flavomarginatus Scirtetellus seminitens Scirtetellus brevipennis Strongylocoris leucocephalus


Continued

4 1

4 2

4 3

4 4

4 5

4 6

4 7

4 8

4 9

5 0

5 1

5 2

5 3

5 4

5 5

5 6

5 7

5 8

5 9

6 0

0 1 1 1 0 1 1 1 1 1

0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 1 1 0

0 1 1 1 1 1 1 1 1 1

0 1 1 1 0 1 1 1 1 0

0 0 0 0 0 0 0 1 1 0

0 ? 0 0 0 0 1 1 1 0

1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1

1 1 1 2 1 2 1 2 2 2

0 0 0 1 0 0 1 1 1 1

0 0 0 0 0 0 0 0 0 0

2 2 1 0 2 2 2 2 2 2

2 1 1 1 1 1 0 2 2 1

0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0

1 0 0 0 0 0 0 0 0 0

0 ? 0 0 1 0 0 1 1 0

1 0 0 0 0 0 0 0 0 0

1 0 0 0 0 0 0 0 0 0

6 1

6 2

6 3

6 4

6 5

6 6

6 7

6 8

6 9

7 0

7 1

7 2

7 3

7 4

7 5

7 6

7 7

7 8

7 9

8 0

0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 1 1 1 1 1 0 0 1 1 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 1 0 1 1 0 0 1

1 1 1 2 0 2 2 2 2 1 0 1 0 2 1 0 0 0 1 1 1 1 0 0 1 1 1 0 0 1 0 0 0 0 0 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1

0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1

0 1 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 1 0 0 1 1 0 0 1 1 0 0 1 0 1 1 1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0

0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0

1 0 1 ? 0 0 0 0 0 0 1 0 0 0 0 0 1 1 0 0 1 0 0 0 0 1 1 1 1 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1

0 0 0 0 0 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

0 1 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0

0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0

1 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 1 0 0 0 1 2 2 2 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 2 2 1 0 0 0 0 0 1 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0

0 1 1 0 0 0 1 0 1 0 1 1 0 1 0 1 0 0 0 0 1 1 1 1 1 1 1 1 0 0 0 0 ? 0 0 0 0 0 1 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 0 0 1 ? 0 0 0 0 0 0 0 0 0 0


658


APPENDIX 2


Continued

8 1

8 2

8 3

8 4

8 5

8 6

8 7

8 8

8 9

9 0

9 1

9 2

? 1 0 ? 0 0 1 1 1 1 1 0 0 0 0 0 1 0 0 1 1 1 0 0 0 1 1 1 1 0 0 1 1 ? 0 1 1 ? 0 0 1 1 1 0

0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 1 1 1 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 ? 0 0 0 0 0 0 0 1 0 0

0 0 1 1 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1 1 0 0 1 0 0 0 0 0

0 0 1 0 0 1 0 1 1 1 1 1 1 1 0 0 0 0 0 0 1 0 1 1 1 0 0 0 0 1 1 1 0 ? 0 0 1 1 1 1 1 0 0 0

0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 1 0 0 0 0 0 0 0 0 0

0 0 1 0 0 1 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 1 1 ? 1 1 1 1 1 1 1 0 0 1

0 0 0 0 1 1 1 1 1 1 1 1 ? 1 1 -

0 0 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 1 1

0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 0 0 1 1 1 1
