Halticini Del Mundo PDF

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 Microtech-
nites 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 dark insects, often with saltatorial hindlegs and

sometimes with ant-like habitus. Many species
The Halticini (Heteroptera: Miridae: Orthotylinae)
exhibit wing polymorphism, with wing reduction in
was first recognized by Costa (1853) as a suprageneric
one or both sexes. All species are thought to be
group of plant bugs. Halticines are generally stout,
phytophagous, with many found on grasses (Schuh &
Slater, 1995). They are mostly distributed in the
Palaearctic zoogeographical region, and are particu-
*Corresponding author. E-mail: n.tatarnic@unsw.edu.au larly species-rich in the Mediterranean as well as
558 © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 558–658
THE HALTICINI OF THE WORLD 559
Eastern Europe and northern Asia (e.g. Mongolia, veins not forming cells; lower portion of the head
Asiatic Russia). Most genera exhibit restricted distri- below eyes greater than twice the height of
butions, perhaps reflecting poor dispersal ability eye. This subtribe is monogeneric, only including
because of the high degree of brachyptery in one or Myrmecophyes comprised of 29 species.
both sexes, which may be attributable to environmen- 2. Halticaria: small, oval, and mostly black; generally
tal stability (Southwood, 1962; Sweet, 1964; Schuh, with both thin, simple setae and pale, scale-like
1974). setae; hemelytra sometimes truncate, in mac-
The taxonomic position of the Halticini within the ropterous females barely covering abdomen, mem-
Orthotylinae has been disputed by several authors. brane when present containing two cells; head
Halticini was defined by Kirkaldy (1902) as a tribe of short and strongly angled backward below the
Orthotylinae, within which he also included the midpoint, eyes overlapping anterolateral margins
Phylinae: Pilophorini (as Pilophorinae sensu Reuter). of pronotum; antennae long and thin, at least as
Wagner (1973) subsequently elevated halticines to long as and often longer than body, with antennal
subfamilial status, segregating them on the basis of segment IV longer than entire hind leg;
male genital structure, thereby recognizing three dis- metafemora generally incrassate; abdomen short
crete suprageneric taxa (Orthotylinae sensu stricto, and rounded, without a narrowly constricted base;
Pilophorinae, and Halticinae). Wagner (1973) further right paramere spoon-shaped with long base and
divided the Halticinae into two tribes, the nomino- a well-defined apical swelling (= apophysis); left
typical Halticini and the Chorosomellini, using char- paramere smaller than right, apically slender,
acters of the head and first antennal segment. angular, with apophysis curved down; phallotheca
Schuh (1974) restored the Halticini as a tribe often black. The Halticaria includes two genera,
within the Orthotylinae, based primarily on the con- Halticus and Barbarosia.
vergent lamellate parempodia. Past polemics have 3. Laboparia: small to medium-sized, either mac-
been recently revisited, with Kiyak (1995) restoring ropterous and elongate-oval or brachypterous and
them to the subfamilial level. Nonetheless, most oval, often strongly sexually dimorphic with male
authors continue to maintain the Halticini as a tribe macropterous and female either brachypterous or
within the Orthotylinae (e.g. Cassis & Gross, 1995; macropterous; membrane with two cells; body with
Kerzhner & Josifov, 1999). both simple setae and scale-like setae; head short,
The aim of this work is threefold: (1) ascertain if angled back ventrally; eyes substylate or stylate;
and how the Halticini are a monophyletic group; antennae with segment III equal to or longer than
(2) determine what taxon relationships a putatively segment IV; abdomen without narrow, ant-like
monophyletic Halticini have; and (3) establish a constriction; legs often short, metafemora some-
generic classification for the Halticini, inclusive of times incrassate; right paramere spoon-shaped,
diagnoses and descriptions. longer than left, usually with apical apophysis; left
paramere thin and hooked, generally with sensory
lobe above base of apophysis.
TAXONOMIC HISTORY
In this classification, the Laboparia are the largest
Costa (1853) diagnosed the halticines on the basis of
subtribe in the Halticini, and encompass all remain-
the tall genae, wide vertex, and saltatorial hindlegs.
ing genera. Schuh (1974, 1975, 1976, 1995) did not
Schuh (1974, 1975, 1976) provided the first phyloge-
accept this subtribal arrangement, and none of the
netic classification of the Miridae, recognizing three
genera described since Wagner (1973) have been sepa-
tribes in the Orthotylinae: Halticini, Nichomachini,
rated into subtribes.
and Orthotylini, all united by the possession of lyre-
Since Costa (1853), genitalic characters have
shaped, apically convergent parempodia and male
played an increasing role in taxonomic delineation in
genitalia with a membranous vesica (= endosoma of
the Miridae (e.g. Reuter, 1883; Knight, 1923; Wagner,
Kerzhner & Konstantinov, 1999), with or without
1940; Southwood, 1953; Kelton, 1955, 1959; Tatarnic
sclerotized spicules.
& Cassis, 2008). In his seminal work on male geni-
Wagner (1973) subdivided the Halticini into three
talia, Kelton (1959) examined nine species from five
subtribes: (1) Myrmecophyaria Reuter, 1891, (2) Hal-
genera of Halticini. He noted that in these taxa, the
ticaria Kirkaldy, 1902, and (3) Laboparia Reuter,
rim of the secondary gonopore is typically oval or
1883, with the following diagnoses:
horseshoe-shaped and the ductus seminis is a flexible
1. Myrmecophyaria; brachypterous morphs ant-like cylindrical tube, both characters shared with the
with a basally constricted and globular abdomen Orthotylini. He also observed that in the Halticini
and very short hemelytra; macropterous morphs both parameres are of similar size and shape, and
without discrete cuneus, hemelytral membrane that except for Halticus (in his work Kelton did not
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 558–658
560 N. J. TATARNIC and G. CASSIS
examine any of the Halticus species that are trans- features characteristic of the Orthotylinae (e.g. lamel-
ferred herein to Microtechnites and Ectmetopterus), late parempodium).
the endosoma possesses several spicules. The definition of the Halticini has remained
The importance of female genitalia in taxonomic unchanged in the nearly 40 years since Schuh (1974),
delineation was first recognized by Slater (1950), who with many character states – in particular those of the
examined four genera of Halticini (Halticus, Labops, female genitalia – unconfirmed for most genera. Addi-
Orthocephalus, and Strongylocoris). Along with iden- tionally, more recent studies indicate another possible
tifying the sclerotized rings of the dorsal labiate plate synapomorphy for the Halticini associated with the
and structures associated with the posterior wall as female vestibulum. When examining the female geni-
useful taxonomic characters, he noted considerable talia of Myrmecophyes oregonensis Schuh & Lattin
variation across taxa examined, leading him to ques- (Schuh & Lattin, 1980), the authors observed that ‘the
tion the validity of contemporary subfamily and tribal small sclerites which lie at the base of the anterior
limits. ovipositor valves [= first gonapophysis] in the vulvar
Kullenberg (1947), Kelton (1959), and Slater area ventral of the bursa copulatrix are symmetrical’,
(1950) all had concerns regarding a classification in marked contrast to the asymmetrical condition
based on pretarsal morphology, believing pretarsal expressed in the remainder of the Orthotylinae (Schuh
structures too labile for establishing higher level & Lattin, 1980). In their examinations of the Halticini
taxonomic relationships. Based on their genitalic species Halticus luteicollis, Pachytomella parallela,
studies, Kelton and Slater both doubted the mono- and an unidentified species of Dimorphocoris, Pluot-
phyly of the Halticini. Slater considered the female Sigwalt & Matocq (2006) also found the sclerites
genitalia to be a more reliable basis for mirid clas- associated with the opening to the vestibulum to be
sification, but for the Halticini found the structure symmetrical, and suggested that this might be the case
of the posterior wall perplexing. Similarly, Kelton for all Halticini. As we show in this work such sym-
believed that male genitalia provided a stable taxo- metry is present in virtually all Halticini with the
nomic character system for the Miridae, but reported exclusion of Goodeniaphila, wherein the left margin of
that within each tribe of the Orthotylinae remark- the vestibular opening has a small accessory sclerite
ably divergent male genitalia were found, implying (Tatarnic, 2009). Symmetry of the vestibular region
non-natural groupings. has also been recently described in other Orthotylini,
Despite these criticisms, Schuh (1974) countered including the Hadronema group and at least one
that along with pretarsal form, characters of the species of Araucanocoris Carvalho, 1983 (Forero,
head and the right paramere are sufficient to group 2008), Mecomma Fieber, 1858, and Harveycapsus
the Halticini, and suggested that the posterior wall dimorpha Cassis, Symonds & Tatarnic (Cassis et al.,
in this tribe may be considered an ‘inherently 2010) (Orthotylini), suggesting that this condition may
variable structure’, which eventually gave rise to prove to be more common than first thought.
the more derived form exhibited in other Orthotyli-
nae. With respect to male genitalic structures, he
argued that although the parameres may be of some
MATERIAL AND METHODS
use in tribal delineations, in general the male geni-
talia do not reveal distinctive trends within the SPECIMENS EXAMINED
Orthotylinae. This study was based on specimens from existing
The first formal tribal description of the Halticini to museum collections and new material collected
include both male and female genitalic characters during the course of the Planetary Biodiversity
was that of Schuh (1974), which included the follow- Inventory study of plant bugs (http://research.amnh.
ing characters: primarily black or dark colouring, org/pbi/index.html). All efforts were made to examine
sometimes with paler markings; body robust to elon- the type species of all genera, and as many species
gate; dorsum generally smooth and impunctate; head within each genus as possible. In the ‘Included
dorsoventrally elongate, with height of gena greater species’ section of each genus treatment, taxa
than the height of the eye; the pronotum and scutel- marked with an asterisk are those that have been
lum not greatly modified except in Myrmecophyes; examined in detail. Diagnoses are provided for all
commonly brachypterous in one or both sexes; metafe- genera of Halticini, whereas redescriptions are
mur frequently incrassate; vesica (= endosoma) mem- included for all genera except for those most recently
branous without spicules; left paramere typically described (Compositocoris, Dampierella, and Good-
elongate and apically hooked; right paramere flat- eniaphila).
tened and apically expanded, generally appearing Specimens were examined from the following col-
club- or spoon-shaped; female posterior wall without lections: AM, Australian Museum, Sydney; AMNH,
inter-ramal lobes; sclerotized rings variable; other American Museum of Natural History, New York;
ANIC, Australian National Insect Collection, CSIRO, TERMINOLOGY

Canberra; BMNH, Natural History Museum, London;
BPBM, Bishop Museum, Honolulu; CNC, Canadian The terminology of morphological structures follows
National Collection of Insects, Agriculture and Agri- modern works such as Schuh (1974) and Cassis &
Food Canada, Ottawa; NHMW, Naturhistorisches Moulds (2002). Terminology for the genitalic struc-
Museum, Vienna; NKUM, Nankai University, Tianjin; tures of both sexes has varied greatly throughout the
QM, Queensland Museum, Brisbane; SAMA, South literature, leading at times to confusion in homology
Australian Museum, Adelaide; SANC, Plant Protec- and terminology. Kerzhner & Konstantinov (1999)
tion Research Institute, Pretoria; TAMU, Texas A&M provided the most recent survey of male genitalic
University, Department of Entomology, College structures, and tabulated structural synonymies. We
Station; TYCN, Tomohide Yasunaga Collection, have taken a more conservative view of the terminol-
Nagasaki; ULB, Université Libre de Bruxelles, Brus- ogy of the aedeagus, and do not follow their subdivi-
sels; USNM, United States National Museum of sion of the endosoma into a proximal conjunctiva and
Natural History, Smithsonian Institution, Washing- distal vesica. We therefore refer to the component of
ton; ZMUC, Danish Natural History Museum, Copen- the aedeagus bounding the secondary gonopore as the
hagen; ZISP, Zoological Institute, Russian Academy of endosoma, based on doubts about the homology of the
Sciences, St. Petersburg. vesica of these authors, as proposed by Cassis,
Schwartz & Moulds (2003) and Cassis (2008). Addi-
tionally, the densely packed sclerotized rings lending
DISSECTIONS AND MORPHOLOGICAL ANALYSIS structural support to the ductus seminis are herein
Genitalia of both sexes were examined by macerating referred to as ‘flexible ribbing’, in order to avoid
the abdomen in 5% KOH at room temperature for confusion with the sclerotized rings of the dorsal
5–10 min. Abdomens were then rinsed in water for labiate plate of the female genitalia.
approximately 10 min before being transferred to Information regarding the female genitalia of the
glycerol for examination, illustration, and storage. Miridae is based on the comparative studies of Slater
For observations of the male aedeagus it was some- (1950) and Davis (1955). The terminology we have
times necessary to remove the phallotheca so that fine used is mostly derived from Davis, but with modifi-
structures of the endosoma could be documented, cation from Cassis et al. (2003) for the posterior wall
particularly for the secondary gonopore and spicules. of the bursa copulatrix, where the K-structures of
As the endosoma is attached to the mesial wall of the Slater (1950) are referred to as the inter-ramal lobes.
phallotheca, it was usually necessary to tear away the These are paired, petiolate, and often spinose sclero-
latter. In other cases, the phallotheca was sufficiently tized lobes, which project into the anterior portion of
translucent to observe all structures without further the bursa copulatrix from the dorsolateral margins
dissection. Dissection of females involved the removal of the inter-ramal sclerites. During the course of this
of ventral abdominal sclerites using forceps and a study we identified similar inter-ramal structures
minuten pin. Subsequently, a minuten was then in several genera of Halticini. In some cases these
dragged between the rami in order to separate the appear as swollen tumescences, whereas in others
posterior wall from the labiate plates. Genitalic illus- they are more akin to the structures characteristic of
trations were created using a camera lucida affixed to the Orthotylini, although never as narrowly basally
a Leica DM LB compound microscope, at 10, 20, and constricted. In this study these are treated as inter-
40¥ power, with 10¥ ocular lenses. Illustrations of ramal lobes, although lacking the basally constricted
parameres are scaled down 50% smaller than aedeagi state expressed in the Orthotylini.
for ease of viewing. In halticines, the sclerotized rings of the dorsal
Gross morphology was observed using a Leica labiate plate (DLP) are often laterally upturned along
MZ12 stereomicroscope. Dorsal habitus photographs with the adjacent margin of the DLP. The DLP also
were taken using the Visionary Digital BK Plus Lab sometimes includes medial sclerotized structures (e.g.
photographic system (http://www.visionarydigital. the inter-ramal bridge in Labops: Slater, 1954). In
com) equipped with a Canon EOS 40D camera. some genera the mesal margins and the rami combine
Because of the small size of many of these insects, with the lateral-most portion of the dorsal labiate plate
multiple shots at different focal depths were taken to form paired medially projecting processes (MP, e.g.
and concatenated using HELICON FOCUS software Anapus americanus, Fig. 10A; Labops, Figs 34G, 35A,
(http://www.heliconsoft.com), and further rendered 36E; Scirtetellus, Fig. 56D). The ventral labiate plate
in Adobe Photoshop CS3. Scanning electron micro- (VLP) is also variable, and is frequently divided, often
graphs (SEMs) of gold coated and uncoated specimens forming a weakly sclerotized band running along the
were taken with a Cambridge scanning electron inner margins of the rami, and medially sometimes in
microscope. the form of a weakly sclerotized plate. Occasionally the
DLP and VLP are closely adpressed in parts, and (excluding uninformative characters), consistency
sometimes together form sclerotized folds or projec- index (CI) = 0.27, retention index (RI) = 0.65, exclud-
tions (e.g. Strongylocoris). ing uninformative characters]. The low CI is indica-
For convenience, the external efferent system of the tive of the high degree of homoplasy exhibited by
metathoracic scent gland is abbreviated as MTG, the multiple characters in the analysis. Nonetheless, the
first labial segment is referred to as LI, and antennal RI of 0.65 indicates a moderately strong phylogenetic
segments are abbreviated as AI through to AIV. 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
PHYLOGENETIC ANALYSIS
[Fig. 2: length (based on equal character weights) =
Ninety-two morphological characters (six uninforma- 448, CI = 0.260, RI = 0.641]. This tree differed some-
tive) were coded for 38 ingroup and five outgroup taxa what from those found under equal weighting,
(see Appendix 1 for the list of characters and charac- although the overall monophyly of the Halticini and
ter states and Appendix 2 for the data matrix). Mul- most subclades were retained. Again, support values
tistate characters were treated as unordered. Rather for most nodes were weak. Subtribal divisions pro-
than coding all species of Halticini we have taken an posed by Wagner (1973) were not supported.
examplar approach to taxon sampling, designed to In their combined morphological and molecular
include the type species of all Halticini genera and analysis of the Cimicomorpha, Schuh, Weirauch &
multiple species from genera where morphological Wheeler (2009) presented a paraphyletic Halticini,
heterogeneity exists (e.g. Anapus, Dimorphocoris, with Halticus and Coridromius forming a sister taxon
Halticus, Microtechnites, Myrmecophyes, Orthoceph- relationship separate from Compositocoris senecionus
alus, and Scirtetellus). Our sampling of Dimorphoc- and Orthocephalus sp., the only other Halticini
oris and Orthocephalus was limited by a lack of included in their analysis. In contrast, our results
availability of specimens. This taxon sampling serves show a monophyletic Halticini, to the exclusion of
as a test of Halticini and halticine generic monophyly Coridromius, which is found instead to be sister to
and sister-taxon relationships, as well as providing a Nichomachus (not included in Schuh et al., 2009).
revised generic framework. Schuh et al. (2009) provide two homoplastic morpho-
Multiple outgroup taxa were selected from the two logical characters – the presence of fleshy apically
other tribes of Orthotylinae: Lattinova jacki Cassis, convergent parempodia and the female posterior wall
an undescribed species of Australian Orthotylus, an lacking K-structures (inter-ramal lobes) – to support
undescribed species of Ceratocapsus, and Aoplonema a Coridromius + Halticus relationship. Both these
princeps (Uhler) were coded from the tribe Orthoty- characters are ubiquitous across the Halticini as well
lini, whereas the Nichomachini was represented by as some outgroup taxa (although see below our dis-
Nichomachus minutus. Resulting trees were rooted cussion of K-structures in Anapus, Labops, and Scir-
with Orthotylus sp. tetellus), and are therefore not informative in this
Parsimony analyses were performed in TNT 1.1, context. Based on our phylogenetic results, we
Willi Hennig Society Edition (Goloboff, Farris & propose a monophyletic Halticini with Coridromius
Nixon, 2008). All characters were treated as unor- removed to form the monogeneric tribe Coridromiini,
dered, with both equal weighting and implied weight- which is sister to the Nichomachini.
ing (Goloboff, 1993) approaches applied to character Below we discuss the major findings of our phylo-
optimization. Tree searching was conducted using the genetic analyses using the implied weighting phylo-
‘traditional’ (i.e. heuristic) search method (max- genetic reconstruction (Fig. 2). As we mapped only
trees = 500 000, starting Wagner trees from random those characters whose reconstruction was unambigu-
seed 0, 5000 replicates, branch swapping by tree ous, some salient traits are not shown on the figure.
bisection-reconnection, ten trees held per replication, This ambiguity is in many cases attributable to our
replacement of existing trees). Implied weights analy- treatment of multistate characters as unordered,
sis used the same settings, with a concavity constant rather than reflective of poor phylogenetic signal
of K = 3. Jackknife values were calculated in TNT for the traits in question. Thus several important
(36% removal probability, with 1000 replications). characters, although not appearing on Figure 2, are
Bremer support values were calculated using the also discussed.
bremer.run script available from the TNT wiki
website (P. Goloboff: http://tnt.insectmuseum.org).
NODE A: NICHOMACHUS + CORIDROMIUS + HALTICINI
PHYLOGENETIC RESULTS
Nichomachus, Coridromius, and the Halticini are
Under maximum parsimony analysis with equal char- united by four unambiguous synapomorphies, three of
acter weighting three trees were found [length = 424 which represent aspects of the female genitalia: the
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 tionally, the lateral margins of the vulvar region (first
(Appendix 1: character 86-1), although these are gonapophyses) are symmetrical (91-1), although in
present in some taxa; the posterior wall lacks inter- our analysis this character is also shared with the
ramal lobes (88-1), although intriguingly these are outgroup species Aoplonema princeps. The lack of
present in Anapus, Labops, and Scirtetellus; and the inter-ramal lobes has previously been used to define
posterior wall lacks a medial region distinctly the Halticini; however here we show inter-ramal lobes
separated from the two lateral regions (90-1). Addi- to be present in three halticine genera, and a lack of
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).
these structures also occurs in Nichomachus and 1), in contrast to the Orthotylini, in which they are
Coridromius; thus their absence is not exclusive to attached basad to the gonopore.
the tribe. One nongenitalic character, incrassate hind Symmetry of the margins of the vestibular opening
femora in females (40-1), is also synapomorphic for (first gonapophyses: 91-1) has recently been proposed
the group. as a potential synapomorphy for the Halticini (Pluot-
Sigwalt & Matocq, 2006). However, although common
to all Halticini, this condition is also exhibited in
NODE B: NICHOMACHUS + CORIDROMIUS several outgroup taxa included in our analysis –
Coridromius, Nichomachus, and Aoplonema princeps
Nichomachus and Coridromius are united by two
(as well as the rest of the Hadronema group: see
male genital characters: the ductus seminis is
Forero, 2008) – and cannot therefore be considered
extremely short and lacks convolutions (58-0) and
synapomorphic for the Halticini.
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
TAXONOMY
group, although in our reconstruction its reconstruc-
tion is ambiguous: only in these two taxa do we see CORIDROMIINI NEW TRIBE
the extreme reduction of the right paramere, which Type genus: Coridromius Montrouzier.
is significantly shorter than the left (50-0). In the
remaining outgroup taxa the parameres are subequal
Type species: Ocypus variegatus Montrouzier, 1861
in length (50-1), whereas across the Halticini the
(monotypy).
parameres are either subequal or the right is much
Ocypus Montrouzier, 1861: 67, junior homonym of
longer than the left (50-2). Paramere size asymmetry
Ocypus Leach, 1819 (Coleoptera).
is well known in other groups of Miridae (e.g.
Coridromius Signoret, 1862: 5 (nom. nov. for
Dicyphina – Cassis, 1986; Termatophylini – Cassis,
Ocypus Montrouzier, 1861).
1995), but is less apparent in the Orthotylinae.
Neocypus Distant, 1914: 378 (unnecessary nom.
nov. for Ocypus Montrouzier, 1861). Carvalho, 1987:
61; Linnavuori, 1994: 15; Schuh, 1995: 46; Cassis &
NODE C: HALTICINI Gross, 1995: 185 [Australian catalogue (cat.); trans-
Characters previously used to diagnose the Halticini ferred to Halticini]; Schuh, 1995: 46 (world cat.; Hal-
– the black coloration of the body (1-1), the saltatorial ticini); Liu & Zhao, 1999: 55; Miyamoto & Yasunaga,
hindlegs (41-1), the apically clubbed or spoon-shaped 1999: 33 (descr.); Chérot, Konstantinov & Yasunaga,
right paramere (49-1), the apically hooked left 2004: 57 (descr.).
paramere (54-1, 2), and the vertically elongate head Coridromoides Carvalho, 1956: 54 (gen. nov.; type
(coded in our analysis as genal length, 13-2, 3) are species: Coridromoides carinatus Carvalho, 1956, by
generally true for the Halticini, although only the last original designation); Tatarnic & Cassis, 2008: 1
of these is synapomorphic for the tribe. In the present (descr.; generic revision).
analysis the rest of these characters exhibit high
degrees of homoplasy (< 0.35 CI). Halticines are also
Diagnosis: Readily distinguished from all other
often recognized by their robust body, although this
Miridae by the following characters: male with hypo-
character is diagnostic for many taxa and clearly
dermic genitalia formed by the coupling of the
exhibits no phylogenetic signal relative to other
left paramere and the aedeagus; female genitalia
Miridae. Only one other unambiguous character is
extremely reduced, with subgenital plate absent,
identified for this node: in macropterous individuals,
vulvar region unsclerotized and symmetrical, poste-
the scutellum is not visible from above (31-0).
rior wall simple and lacking medial area and inter-
However, visibility of the mesoscutum is variable
ramal lobes. Additionally, body short and squat,
above and below this node, and we do not consider it
metafemora greatly swollen and modified for
a particularly informative or meaningful trait.
jumping, hemelytra strongly deflected at cuneus.
Based on limited sampling Schuh (1974: table 3)
proposed that the Halticini do not possess endosomal
spicules. Here we show that endosomal spicules are Diversity and distribution: Coridromius currently
actually present in most halticines (72-1), although includes 33 species found throughout the Old World
occasionally absent (spicules are lacking in Halticus, tropics and subtropics, with the Australian species
Namaquacapsus, Piezocranum and Plagiotylus). Coridromius chenopoderis Tatarnic & Cassis intro-
When present the endosomal spicules of the Halticini duced to Hawaii, south-western USA, and northern
are positioned distad to the secondary gonopore (73- Mexico (Tatarnic & Cassis, 2008).
Remarks: In our analysis Coridromius does not fall Dicyphopsis Poppius, 1914 [4] Ethiopian region
within the Halticini. This is unsurprising, as it lacks Dimorphocoris Reuter, 1890 [52] Palaearctic and
many of the characteristics typical of the tribe: elon- African region
gate bases to both parameres, a spoon-shaped right Ectmetopterus Reuter, 1906 [6] Palaearctic and
paramere, mostly black coloration, the aedeagus with Oriental region
a well-developed endosoma (but see Halticus), the Euryopicoris Reuter, 1875b [2] Palaearctic region
ductus seminis with sclerotized ribbing and sclero- Goodeniaphila Tatarnic [2] Australia
tized secondary gonopore, and well-developed sclero- Halticus Hahn, 1832 [18] Eastern Hemisphere
tized rings with upturned lateral margins. Indeed, and North America
most genitalic characters in Coridromius are unlike Labops Burmeister, 1835 [12] Holarctic
those of any Halticini (or any other mirid): the female Microtechnites Berg, 1879 [5] reinstated
genitalia are highly reduced with the sclerotized rings status Neotropical and Nearctic regions
completely absent, and in the male the aedeagus and Myrmecophyes Fieber, 1870 [27] Holarctic
left paramere are coupled to form a piercing intromit- Namaquacapsus Schuh, 1974 [1] Ethiopian region
tent organ. Nanniella Reuter, 1904 [6] Ethiopian region
Coridromius is the only plant bug genus known to Orthocephalus Fieber, 1858 [23] Palaearctic and
practice traumatic insemination, whereby males use Nearctic regions
hypodermic genitalia to stab females in the abdomen Pachytomella Reuter, 1890 [7] Palaearctic region
during mating, completely bypassing the female Piezocranum Horváth, 1877 [4] Palaearctic region
genital tract (Tatarnic et al., 2006). As a result, Plagiotylus Scott, 1874 [7] Palaearctic region
females have extremely reduced genitalia, with some Schoenocoris Costa, 1842 [1] Palaearctic region
species exhibiting various paragenital modifications Scirtetellus Reuter, 1890 [14] Palaearctic region
at the site of insemination, believed to reduce mating- Strongylocoris Blanchard, 1840 [16] Palaearctic
induced costs (Tatarnic et al., 2006, Tatarnic & Cassis region
2010). Because of this, the use of genitalic traits – in
particular genital reduction or simplification – should
not be taken as evidence of common ancestry. HALTICINI COSTA (FIGS 3–58)
We believe that Coridromius is sufficiently different Halticini Costa, 1853: 75 (new tribe); Kirkaldy, 1906:
from all other Halticini (as well as all other existing 130 (cat.); Knight, 1923: 497, 498 (gen. key); Blatch-
tribes within the Orthotylinae) to justify the erection ley, 1926: 797 (east US gen. key); Hsiao, 1942: 253
of its own monotypic tribe. Ultimately its relationship (Chinese gen. key); Wagner, 1952: 95 (desc.; Palaearc-
to other tribes of Orthotylinae should not be deter- tic gen. key); Carvalho & Leston, 1952: 245 (British
mined through morphology alone, but will instead gen. key); Carvalho, 1952: 39, 40, 73 (list); Carvalho,
necessitate inclusion of other data (i.e. DNA). 1955: 65 (world gen. key); Carvalho, 1958: 5 (cat.);
A thorough revision of the genus can be found in Slater, 1950: 43 (female genitalia); Kelton, 1959: 33
Tatarnic & Cassis (2008). (male genitalia); Wagner, 1961: 47 (European fauna);
Wagner & Weber, 1964: 250 (French fauna); Wagner,
1973: 2 (Mediterranean fauna); Schuh, 1974: 26, 273
HALTICINI (classification); Schuh, 1975: 13 (femoral trichoboth-
(Numbers following generic names indicate numbers ria); Schuh, 1976: 18, 35 (pretarsus; classification);
of included species.) Linnavuori, 1994: 4 (West African gen. key); Schuh &
Slater, 1995: 44 (world cat.); Kerzhner, 1988: 778, 826
(key to gen.; Palaearctic fauna); Kerzhner & Konstan-
CHECKLIST OF GENERA OF HALTICINI tinov, 1999: 124 (male genitalia).
Acratheus Distant, 1910 [3 species] Oriental and Halticocoridae Douglas & Scott, 1865: 35 (fam.
African region stat.).
Anapus Stål, 1858 Stiphrosomatidae Douglas & Scott, 1865: 35 (new
= Platyporus Reuter, 1890; new synonymy fam.).
[8] Holarctic Halticaria Kirkaldy, 1902: 139 (list); Reuter, 1910:
Barbarosia Kiyak, 1995 [2] Palaearctic region 115 (desc.; key); Poppius, 1911: 33 (class.); Poppius,
Chorosomella Horváth, 1906 [2] Palaearctic 1914: 59, 82 (key; descr.); Van Duzee, 1916: 43 (cat.);
region Butler, 1923: 463 (descr.; key); Wagner, 1973: 15
Compositocoris Schwartz, Schuh & Tatarnic (European fauna).
[1] Ethiopian Region Diplacaria Reuter, 1883: 564: (new division).
Dampierella Tatarnic, 2009 [1] Australia Laboparia Reuter, 1883: 567 (new division; gen.
Dasyscytus Fieber, 1864 [1] Palaearctic region key); Puton, 1886: 53 (cat.); Atkinson, 1890: 117 (cat.);
KEY TO THE GENERA OF HALTICINI

1. 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
2. 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
3. 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
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
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
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
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
8. 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
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
10. 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
11. 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
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
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
14. 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
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.); mostly light brown, tan or green; body generally stout
Reuter, 1905a: 19, 28 (descr.; key); Reuter, 1909: 71 and compact, sometimes elongate; hindlegs often sal-
(list); Oshanin, 1910: 793 (Palaearctic cat.); Oshanin, tatorial; pronotal texture most often smooth and
1912: 73 (Palaearctic cat.); Van Duzee, 1916: 211, 373 glossy, sometimes punctate; vestiture variable, either
(key; descr.); Wagner, 1973: 15 (subtribe; European only with thin simple setae, or also with pale and
fauna). lamellate setae; legs and antennae often spinose;
Labopini Knight, 1923: 501 (key); Blatchley, 1926: head typically dorsoventrally elongate, height of
797 (east US gen. key); Hedicke, 1935: 53 (gen. key); genae almost always equal to or greater than height
Knight, 1941: 8, 19, 74 (Illinois gen. key); Kiritsh- of eye; pronotum campanulate, rectangulate or trap-
enko, 1951: 117 (USSR gen. key). ezoidal; aedeagus with thinly sclerotized phallotheca,
Myrmecophyaria Reuter, 1891: 106 (new division); apex of phallotheca most often constricted, sometimes
Wagner, 1973: 15 (subtribe; European fauna). keeled, rarely with thin apical projections, ductus
Halticarini Zimmerman, 1948: 198 (key). seminis long or short with flexible ribbing formed by
numerous thin, closely packed sclerotized rings,
Diagnosis: Usually black or dark brown with or sometimes with elongate subapical segment lacking
without lighter contrasting markings, sometimes flexible ribbing; secondary gonopore weakly to
Figure 3. Photographs of Halticini genera: Acratheus–Halticus. Abbreviations: F, female; M, male.
Figure 4. Photographs of Halticini genera: Labops–Strongylocoris. Abbreviations: F, female; M, male.
strongly sclerotized, often with distinctive scale-like point of eye; elongate, narrow, AI only slightly thicker
texturing, endosoma almost always present as a than AII. Labium (Fig. 5B): extending beyond meta-
membranous bag within the phallotheca (except in coxae, LI somewhat swollen and elongate; thorax
Halticus), often with one or several sclerites, some- (Figs 3, 5B, D–F): pronotum trapezoidal, steeply
times with fields of spines or dentitions; basal portion inclined, collar flat, visible from above, posthumeral
of both parameres usually elongate (except in angles distinctly depressed, posterior margin cari-
Dampierella and Goodeniaphila, where the base of nate, medially concave; mesoscutum visible; scutel-
the left paramere is relatively short with respect to lum medially raised, basally depressed, laterally
other halticines), both parameres ventrally concave; declivent; metathoracic spiracle exposed, operculum
left paramere usually L-shaped, sometimes with large, without evaporative bodies; MTG external
swollen sensory lobe, with long apophysis, often api- efferent system reduced, tongue-like, peritreme
cally hooked and sometimes bifid; right paramere located along posterior margin of metepisternum,
flattened and generally spoon- or club-shaped, some- narrow strip of evaporative areas anteriad to peri-
times with a small apical apophysis; posterior wall of treme. Hemelytra (Figs 3, 5A): Laterally declivent at
female without inter-ramal lobes (but see Anapus, corial fracture, lateral margin slightly flared apically;
Scirtetellus, and Labops); sclerotized rings variable, membrane extending beyond abdomen. Legs (Figs 3,
but generally weakly concave or laterally upturned; 5G): elongate, narrow; metafemora not incrassate;
margins of first gonapophyses symmetrical, some- pretarsi without pulvilli. Abdomen: elongate-ovoid.
times with weak sclerotization. Male genitalia (Figs 6A–C, 5H): pygophore conical,
simple, without modifications or processes; parameres
subequal in length, both with elongate sensory lobe;
ACRATHEUS DISTANT (FIGS 3, 5–6)
left paramere broad, with thin, strongly arcuate distal
Acratheus Distant, 1910a: 16 (gen. nov. type species: arm; right paramere spoon-shaped with thin apical
Acratheus nocturnus Distant, 1910 by monotypy); apophysis, projecting beyond genital opening of pygo-
Distant, 1910b: 284 (India); Reuter, 1910: 161 (cat.); phore; phallotheca simple, elongate, abruptly narrow-
Carvalho, 1952: 73 (list); Carvalho, 1955: 67 (key); ing from midpoint to apex; ductus seminis short, with
Carvalho, 1958: 6 (cat.), Linnavuori, 1994: 7 [diagno- irregular U-shaped sclerotized secondary gonopore,
sis (diag.), key to spp.]; Schuh, 1995: 45 (cat.) opening into a conical sclerotized tube running length
of phallotheca. Female genitalia (Fig. 6D, E): sclero-
Diagnosis: Acratheus is distinguished from other Hal- tized rings minute, thin, lateral margins and adjacent
ticini except Nanniella by the densely punctate portion of DLP weakly upturned; posterior wall mem-
scutellum, pronotum, and thoracic pleura. It differs branous, with very weak lateral swellings; margins of
from Nanniella by the lack of deep punctures on the vestibulum symmetrical with medial margins of first
frons and vertex and the shape of the left paramere, gonapophyses forming slightly tumescent, weakly
which in Nanniella is more elongate and apically sclerotized plates.
barbed.
Redescription: Macropterous, body length 2–3 mm. Diversity and distribution: Acratheus has four
Coloration (Fig. 3): mostly dark brown to black, with species, two in India and two in equatorial Africa.
yellow or yellow-brown markings. Surface and vesti- Lack of records despite intensive sampling in Middle
ture (Figs 3, 5A–H): head mostly smooth with shallow Asia by Linnavuori (1975, 1984, 1986, 1992) suggests
punctures limited to genae (Fig. 5A–C). Pronotum, this apparent geographical disjunction is real.
scutellum, and thoracic pleura densely punctate
(Figs 3, 5A–C, E–F) Hemelytra densely punctate, less Included species: Acratheus albipes (Motschulsky,
so towards apex of corium (Figs 3, 5A); cuneus punc- 1863) India
tate. Abdominal sternites with few punctures later- Acratheus nocturnus Distant, 1910 India
ally (Fig. 5H). Thorax and abdomen sparsely covered Acratheus ocellaris Linnavuori, 1994* Benin;
with longish, brown, decumbent setae (Fig. 5B, E, H). Ivory Coast; Nigeria; Togo
Structure: body elongate and slightly ovate. Head Acratheus punctiger (Linnavuori, 1975)* Benin;
(Figs 3, 5A–C): transverse, short, taller than wide; Nigeria; Sudan; Upper Volta
eyes large, height of gena slightly greater than eye
height; posterior margin of vertex carinate, sublinear
with eyes not touching pronotum; vertex convex; Biology and host plant associations: Acratheus is
clypeus not distinctly projecting forward; maxillary found on undergrowth in savannah and rain forests
plate long and vertical; bucculae small, narrow. (Linnavuori, 1994). No host plants have been
Antennae (Figs 3, 5A–C): insertion in line with mid- recorded for this genus.
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 separate evolutionary lineage’, and posited that Acra-
genera characterized by deep punctations on the theus and Halticus are closely related, based on the
head, pronotum, thoracic pleura, and hemelyra, the following shared similarities: an elongate head,
other being the endemic African genus Nanniella. swollen first labial segment (LI), and halticine-type
Schuh (1974) first proposed a close relationship parameres. He also noted that in Nanniella the head
between these genera. In contrast, Linnavuori (1994) is notably shorter and LI is only slightly thicker than
stressed that Nanniella ‘undoubtedly represents a LII.
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 described the parameres of the former species, which
results (Figs 1, 2), we concur with Schuh (1974) that are very similar to the African species.
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 ANAPUS STÅL (FIGS 3, 7–10)
first labial segment is only slightly less thick in Anapus Stål, 1858: 188 (gen. nov.; type species:
Nanniella than in Acratheus. Conversely, both Anapus kirshbaumi Stål, 1858: 189 by monotypy);
genera share many characters, with the following Walker, 1873: 160 (list); Reuter, 1891: 69, 159 (key;
significant similarities: the posterior margin of the descr.); Kirkaldy, 1906: 131 (list); Hueber, 1906: 5
pronotum is concave; the configuration of the exter- (key); Oshanin, 1910: 787 (cat.); Reuter, 1910: 147
nal efferent system of the metathoracic gland; the (cat.); Stichel, 1933: 235 (key); Kiritshenko, 1951: 126
secondary gonopore is wide and distally modified (key); Wagner, 1952: 96, 106 (key; descr.); Carvalho,
into a sclerotized tube running the length of the 1955: 66 (key); Carvalho, 1958: 6 (cat.); Kerzhner,
phallotheca; the shape of the parameres; the poste- 1964a: 965 (diag., key to spp.); Wagner, 1973: 55
rior wall of the bursa copulatrix is membranous to (descr., key to spp.); Schuh, 1995: 45 (world cat.).
weakly sclerotized, bearing two minor lateral swell- Labops (Merotrichaea) Reuter, 1875c: 24 (subgen.
ings; and the sclerotized rings of the female dorsal nov.; type species: Orthocephalus freyi Fieber, 1864,
labiate plate are weakly sclerotized and upturned by subsequent designation Kirkaldy, 1906); Reuter,
laterally. 1875b: 87 (key); Reuter, 1891: 79 (synonymy).
This redescription of Acratheus is based on exami- Merotrichia Carvalho, 1958: 6 (cat.; generic status).
nation of the African species (Ac. ocellaris and Platyporus Reuter, 1890: 246 (gen. nov.; type
Ac. punctiger) and the literature (Linnavuori, 1994). species: Platyporus dorsalis Reuter, 1890 by mono-
We did not have access to the Indian species Ac. al- typy); Carvalho, 1958: 30 (cat.); Wagner, 1973: 53
bipes and Ac. nocturnus; however, Linnavuori (1994) (descr.); Schuh, 1995: 69 (world cat.). New synonymy.
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 trally, with prised operculum; and, female posterior
characters: substylate eyes; antennal insertion below wall of bursa copulatrix often with bilaterally sym-
ventral margin of eye; AI swollen and longer than metrical, sclerotized inter-ramal tumescences.
height of eye, with several elongate spines; hemelytra
of brachypterous individuals very short; male ductus Redescription: Coloration (Fig. 3): Body uniformly
seminis long, with elongate apical section lacking black with minor yellow and orange markings;
ribbing; secondary gonopore rounded, opening ven- appendages black. Surface and vestiture (Figs 3,
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, on metatibiae. Structure: both sexes usually brac-
scutellum and hemelytra rugulose, often with hypterous, rarely macropterous. Head (Figs 3, 7A–C,
wrinkles on vertex and frons radiating from midline. E): transverse, triangular in shape in dorsal view,
Body with erect and semi-erect simple setae, some- with posterior margin straight to very weakly
times intermixed with dense decumbent, white, scale- concave; head in lateral view angled somewhat cau-
like setae. Head with a few elongate thin erect black dally below antennae; height of head below eye
setae. Antennae with semi-erect, spine-like setae; AI approximately twice height of eye; clypeus strongly
with a few to numerous elongate spines, especially in declivent, not projecting forward; posterior margin of
males. Legs with elongate sharp spines, particularly vertex not carinate; frons rounded, strongly declivent
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 margins rounded; posterior margin weakly concave;
anterior of pronotum; eyes substylate; Antennae mesoscutum only visible in macropterous individuals;
(Figs 3, 7A–C, E): antennal insertion well below eye; scutellum triangular, flat to tumescent; metathoracic
subequal to body length; AI thickened, sometimes spiracle large and exposed, sometimes with evapora-
incrassate; remaining segments thin. Labium tive bodies along posterior margin; MTG external
(Fig. 7E): length variable, reaching from mesocoxae to efferent system broad, swollen and triangular, gener-
basal abdomen sternites; LI incrassate. Thorax ally reaching lower margin of metathoracic spiracle;
(Figs 3, 7A–D): pronotum short, weakly declivent in peritreme oval, medially on metepisternum, subpar-
lateral view, rectangular in brachypterous forms, allel to posterior margin of pleuron. Hemelytra
trapezoidal in male macropterous individuals; collar (Figs 3, 7A–B): usually brachypterous, sometimes
absent; callosite region sometimes distinct, lateral reduced to a small pad, in macropterous morphs with
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 along with adjacent portion of DLP, DLP sometimes
(Figs 3, 7A–B, E–F): metafemora strongly incrassate; with heavily sclerotized inter-ramal bridge; margin of
foretibiae usually swollen apically and weakly VLP adjacent to rami sclerotized, medially with a
arcuate; pretarsi with elongate fleshy parempodia small flange; posterior wall of bursa copulatrix
(except Anapus americanus). Abdomen (Figs 3, 7G): variable, but always with bilaterally symmetrical,
pear-shaped, elongate oval, or oval. Male genitalia sclerotized structures (inter-ramal tumescences),
(Figs 8A–C, 7B, E, G, 9A–C): pygophore triangular, sometimes toothed or serrate; margins of vestibular
posterior margin concave at left paramere insertion; opening symmetrical with weak sclerotization.
right paramere sometimes projecting slightly from
pygophore; phallotheca simple, sometimes moderately
Diversity and distribution: Anapus has seven Holarc-
sclerotized apically; left paramere sensory lobe broad,
tic species, one from the western USA and the
apophysis thin, hooked; right paramere broad, spoon-
remainder Palaearctic.
shaped, with small apical apophysis; phallotheca
simple, without processes; ductus seminis elongate
with flexible ribbing, sometimes very slender (as in Included species: Anapus americanus Knight,
Anapus freyi), subapically with elongate section, 1959* USA
sometimes heavily sclerotized, lacking flexible Anapus dorsalis (Reuter, 1890)* comb. nov. Rus-
ribbing; secondary gonopore elongate, U-shaped, with sia; Turkey
semiclosed operculum; endosoma with one or two Anapus freyi (Fieber, 1864)* Russia
simple spicules of varying size and shape, in most = Pachytoma jakovloeffi Reuter, 1879 (syn. Reuter,
species also with paired, elongate, and sclerotized 1881)
spiculate strips near apex. Female genitalia (Figs 8D, Anapus kirschbaumi Stål, 1858* Russia
E, 10A–C, 10A–F): sclerotized rings moderate to = Orthocephalus opacus Jakovlev, 1875 (syn. Jak-
large, elongate, lateral margins weakly upturned ovlev, 1877)
= Anapus pectoralis Horváth, 1904 (syn. Kerzhner, inter-ramal lobes of the posterior wall (characters 82
1996) and 88) are both shared with Labops (Slater, 1950;
Anapus longicornis Jakovlev, 1882* Hungary; Slater, 1954), with the latter of these structures also
Russia found in Scirtetellus. In these other genera and
= Labops (Pachytoma) arenarius Horváth, 1884 An. americanus, the inter-ramal lobes are hollow, api-
(syn. Reuter, 1890) cally tapered, and medially convergent, and spinose.
= Anapus flavicornis Reuter, 1904 (syn. Kerzhner, These appear homologous to the inter-ramal lobes
1970) found in the Orthotylini (Slater, 1950; also known as
Anapus pachymerus (Reuter, 1881) Spain K-structures), although they differ in that they are
Anapus rugicollis (Jakovlev, 1877)* Russia generally broader and much less narrowly constricted
= Pachytoma sibirica Sahlberg, 1878 (syn. Kerzh- basally (89-1).
ner, 1962) Aside from An. pachymerus, all species of Anapus
= Pachytoma nigrita Jakovlev, 1881 (syn. Kerzhner, were examined for this study.
1962)
BARBAROSIA KIYAK, 1995 (FIGS 3, 11–12)
Biology and host plant associations: Host plant
records are known for four species of Anapus (Table 1; Halticidea Reuter, 1901: 172 (gen. nov.; junior
Schuh, 1995). Anapus freyi is known from a grass homonym of Halticidea Hohrn, 1893 [Coleoptera];
species, and the other three species are known from type species: Halticidea punctulata Reuter, 1901 by
either euasterid or eurosid angiosperms. monotypy); Kirkaldy, 1906: 130 (list); Oshanin, 1910:
802 (Palaearctic cat.); Reuter, 1910: 147 (cat.); Kirit-
Remarks: Herein we propose Platyporus as a junior shenko, 1951: 127 (key); Carvalho, 1952: 73 (list);
synonym of Anapus, resulting in the new combina- Carvalho, 1955: 68 (key); Carvalho, 1958: 11 (cat.);
tion: Anapus dorsalis (Reuter, 1890). This species Wagner, 1973: 13 (descr.; key); Schuh, 1995: 53 (world
conforms to the generic diagnosis of Anapus given in cat.).
this work, and shares significant overlap in salient Barbarosia Kiyak, 1995: 216 (nom. nov. for Halti-
morphology, and striking similarities in the male cidea Reuter, 1901).
aedeagus and female posterior wall of the bursa copu-
latrix. This synonymy is supported by our phyloge- Diagnosis: Barbarosia is recognized by the following
netic analysis. combination of characters: coleopteroid; mostly shiny
In all the Old World species of Anapus examined black; white scale-like setae on laterotergites; frons
the male genitalia are nearly identical, with the fol- extending beyond clypeus; eyes substylate; antennae
lowing similarities: the distal portion of the ductus short, subequal to body length; left paramere
seminis is sclerotized, the endosoma is relatively L-shaped; endosoma with single S-shaped spicule;
small, bearing one or two spicules and apically with and, female DLP exaggerated, sclerotized ring
sclerotized fields of small spines, and the left strongly convoluted. It is similar to Halticus but with
paramere has a broad sensory lobe. However, in the shorter antennae, which in the male are slightly
New World species, An. americanus, the distal portion longer, and in the female slightly shorter than the
of the ductus seminis is weakly sclerotized, the endo- body. The female sclerotized rings are unlike those of
soma is much larger with the apical fields of small any other halticine.
spines only faintly visible, and the left paramere lacks
the large sensory lobe present in other species. Addi- Redescription: Coleopteroid, 2.3–3.35 mm, male
tionally the body is much more ant-like in shape, the slightly smaller than female. Coloration (Fig. 3): body
hemelytra are reduced to very small pads, the pre- mostly shining dark brown to black with paler mark-
tarsi lack fleshy pulvilli, and the first antennal ings. Surface and vestiture (Figs 3, 11A–E): body
segment is thinner, more elongate, and has fewer sparsely clothed in pale decumbent simple setae and
black spines. Despite these differences we consider slightly thicker, white scale-like setae on lateroterg-
An. americanus to represent a divergent New World ites. Antennae with semidecumbent, spine-like setae,
lineage within Anapus rather than a new monotypic AI with several black spines. Legs with black spines.
genus, based primarily on commonalities in the struc- Frons (both sexes) and AI (males) sometimes with
ture of the female posterior wall of the bursa copula- long white setae (punctulata). Head mostly smooth,
trix. Again, this is supported by our phylogenetic pronotum rugulopunctate, scutellum rugulose, hem-
results. elytra finely punctate. Structure: head (Figs 3, 11A–
Anapus shares some notable female genitalic pecu- C): Short, transverse; vertex broad, posterior margin
liarities with several other taxa: in particular, the not carinate; frons bulbous, extending beyond
sclerotized inter-ramal bridge of the DLP and the clypeus; genae tall, approximately twice height of eye;
Table 1. Host plant records for the Halticini
Genus Species Host plant Family Reference
Anapomella
arnoldii Helichrysum arenarium Asteraceae Kerzhner, 1964a
arnoldii Helichrysum arenarium Asteraceae Putshkov, 1961
Anapus
dorsalis Alyssum calycinum Brassicaceae Seidenstücker, 1959
freyi Agropyron sp. Poaceae Kerzhner, 1964a
kirshbaumi Artemisia sp. (Seriphidium) Asteraceae Kerzhner, 1964a
longicornis unknown sp. Apiaceae Kerzhner, 1964a
rugicollis Medicago sp. Fabaceae Kerzhner, 1964a
Barbarosia
punctulata Elymus junceus Poaceae Kerzhner, 1964a
Chorosomella
jakowleffi unknown sp. Poaceae Kerzhner, 1964a
jakowleffi unknown sp. Poaceae Wagner, 1973
jakowleffi Crimea sp. Poaceae Wagner, 1973
Compositocoris
senecionus Senecio laxus Asteraceae Schwartz et al., 2008
Leysera tenella Asteraceae Schwartz et al., 2008
Ursinia sp. Asteraceae Schwartz et al., 2008
Dampierella
schwartzi Dampiera incana incana Goodeniaceae Tatarnic, 2009
Dasyscytus
sordidus Rhanterium epapposum Asteraceae Linnavuori, 1986
Dimorphocoris
bleusi Stipa tenacissima Poaceae Wagner, 1973
cilix unknown sp. Poaceae Wagner, 1973
lateralis unknown sp. Poaceae Wagner, 1973
lurensis unknown sp. Poaceae Wagner, 1973
osellai Lotus sp. Fabaceae Tamanini, 1975
osellai Trifolium sp. Fabaceae Tamanini, 1975
pygmaeus unknown sp. Poaceae Wagner, 1973
schmidti unknown sp. Poaceae Franz & Wagner, 1961
seidenstueckeri unknown sp. Asteraceae Linnavuori, 1984
seidenstueckeri Teucrium sp. Lamiaceae Linnavuori, 1984
seidenstueckeri Trifolium sp. Fabaceae Linnavuori, 1984
tauricus Festuca beckeri Poaceae Kerzhner, 1964b
tauricus Festuca sulcata Poaceae Kerzhner, 1964b
tauricus Festuca taurica Poaceae Kerzhner, 1964b
tauricus Poa sp. Poaceae Kerzhner, 1964b
tomasi Festuca sp. Poaceae Tamanini, 1975
tomasi Stipa sp. Poaceae Tamanini, 1971
tomasi Stipa sp. Poaceae Tamanini, 1976
tristis Anthemis sp. Asteraceae Linnavuori, 1992
Euryopicoris
nitidus unknown sp. Poaceae Wagner, 1973
Goodeniaphila
Cassisi Coopernookia strophiolata Goodeniaceae Tatarnic, 2009
Goodeniaphila
schuhi Goodenia amplexans Goodeniaceae Tatarnic, 2009
Scaevola basedowii Goodeniaceae Tatarnic, 2009
Scaevola ovalifolia Goodeniaceae Tatarnic, 2009
Halticus
apterus Vicia striata Fabaceae Hoberlandt, 1956
apterus Vicia sp. Fabaceae Wagner, 1973
apterus Ononis sp. Fabaceae Wagner, 1973
beganus Phaseolus vulgaris Fabaceae Alayo, 1974
beganus ‘rotten oak’ Fagaceae Henry & Smith, 1979
insularis cucumber Cucurbitaceae Carvalho, 1956
insularis Ipomoea sp. Convolvulaceae Carvalho, 1956
intermedius Anemone canadensis Ranunculaceae Knight, 1968
intermedius Clematis ligusticifolia Ranunculaceae Knight, 1968
luteicollis unknown sp. Poaceae Ehanno, 1960
luteicollis unknown sp. Fabaceae Josifov, 1974
luteicollis Clematis vitalba Ranunculaceae Wagner, 1973
macrocephalus Carduncellus sp. Asteraceae Wagner, 1973
macrocephalus Clematis sp. Ranunculaceae Wagner, 1973
pusillus Ononis sp. Fabaceae Göllner-Scheiding, 1972
pusillus Galium verum Rubiaceae Wagner, 1973
saltator cucumber Cucurbitaceae Wagner, 1973
saltator Calendula sp. Asteraceae Wagner, 1973
saltator Althaea sp. Malvaceae Wagner, 1973
tibialis Ipomoea sp. Convolvulaceae Carvalho, 1956
tibialis Phaseolus sp. Fabaceae Carvalho, 1956
tibialis unknown sp. Cucurbitaceae Carvalho, 1956
Labops
hesperius Agropyron cristatum Poaceae Kelton, 1980
hesperius Rosa arkansana Rosaceae Kelton, 1980
hesperius unknown sp. Poaceae Fuxa & Kamm, 1976
hesperius Koeleria cristata Poaceae Mills, 1939
hesperius Poa secunda Poaceae Mills, 1939
hesperius Stipa comata Poaceae Mills, 1939
hesperius Stipa williamsi Poaceae Mills, 1939
hesperius Hordeum sp. Poaceae Mills, 1939
hesperius Triticum sativum Poaceae Mills, 1939
hesperius wheat (Triticum sp.) Poaceae Mills, 1939
hirtus wheat (Triticum sp.) Poaceae Mills, 1939
hirtus wheat (Triticum sp.) Poaceae Mills, 1939
Table 1. Continued
Genus Species Host plant Family Reference
Microtechnites
bractatus Bryonia sp. Cucurbitaceae Linnavuori, 1984
bractatus Cucurbita moschata Cucurbitaceae Maldonado, 1969
bractatus Ipomoea batatas Convolvulaceae Maldonado, 1969
bractatus Fraxinus sp. Oleaceae Linnavuori, 1984
Myrmecophyes
alboornatus ‘native spp.’ Poaceae Bykov, 1971
alboornatus Agropyron sp. Poaceae Kerzhner & Yachevskii, 1967
frontosus Psathyrostachys juncea Poaceae Drapolyuk & Kerzhner, 2000
macrotrichus Festuca sp. Poaceae Bykov, 1971
macrotrichus Carex sp. Cyperaceae Bykov, 1971
oregonensis Agropyron spiculatum Poaceae Schuh & Lattin, 1980
oregonensis Festuca idahoensis Poaceae Schuh & Lattin, 1980
trispiculus Aremisia terrae-albae Asteraceae Drapolyuk & Kerzhner, 2000
Orthocephalus
bivittatus Artemisia sp. Asteraceae Kerzhner, 1964a
brevis Campanula rapunculoides Campanulaceae Göllner-Scheiding, 1972
brevis Campanula rapunculoides Campanulaceae Wagner, 1973
brevis Campanula sp. Campanulaceae Hoberlandt, 1963
coriaceus Quercus pedunculata Fagaceae Ehanno, 1965
coriaceus Quercus sp. Fagaceae Ehanno, 1960
coriaceus not specified Asteraceae Göllner-Scheiding, 1972
coriaceus Tanacetum sp. Asteraceae Wagner, 1973
coriaceus Achillea sp. Asteraceae Wagner, 1973
coriaceus Centaurea sp. Asteraceae Wagner, 1973
funestrus Artemisia gigantea Asteraceae Kerzhner, 1978
funestrus Artemesia stelleriana Asteraceae Kerzhner, 1978
medvedvi Serratula sp. Asteraceae Kerzhner, 1964a
melas Centaurea sp. Asteraceae Seidenstücker, 1962
melas Centaurea sp. Asteraceae Wagner, 1973
minimus near Cousinia or Scorzonera Asteraceae Drapolyuk & Kerzhner, 2000
proserpinae Carduus spp. Asteraceae Wagner, 1973
proserpinae Scolymus spp. Asteraceae Wagner, 1973
proserpinae Silybum spp. Asteraceae Wagner, 1973
saltator not specified Poaceae Ehanno, 1960
saltator not specified Asteraceae Göllner-Scheiding, 1972
saltator not specified Poaceae Gravestein, 1978
saltator not specified Asteraceae Hoberlandt, 1956
scorzonerae Scorzonera tau-saghyz Asteraceae Drapolyuk & Kerzhner, 2000
scorzonerae Takhtajaniantha pusilla Asteraceae Drapolyuk & Kerzhner, 2000
vittipennis Achillea sp. Asteraceae Göllner-Scheiding, 1972
vittipennis Anthemidae sp. Asteraceae Hoberlandt, 1956
vittipennis Chrysanthemum sp. Asteraceae Göllner-Scheiding, 1972
vittipennis Ononis spinosa Fabaceae Göllner-Scheiding, 1972
vittipennis Chrysanthemum leucanthemum Asteraceae Wagner, 1973
Pachytomella
doriae Plantago albicans Plantaginaceae Wagner, 1973
parallela Potentilla sp. Rosaceae Wagner, 1973
parallela Potentilla sylvestris Rosaceae Kerzhner, 1964A
passerinii Thapsia garganica Apiaceae Wagner, 1973
Plagiotylus
maculatus Teucrium chamaedrys Lamiaceae Ribes, 1984
maculatus Teucrium chamaedrys Lamiaceae Wagner, 1973
ruffoi Astragalus siculus nebrodensis Fabaceae Tamanini, 1960
ruffoi Prangos sp. Apiaceae Tamanini, 1960
ruffoi Astragalus siculus nebrodensis Fabaceae Wagner, 1973
zorzii Globularia sp. Globulariaceae Wagner, 1973
Schoenocoris
flavomarginatus not specified Juncaceae Wagner, 1973
Strongylocoris
atrocoeruleus Peucedanum oficinale Apiaceae Wagner, 1973
niger Peucedanum oreoselinum Apiaceae Wagner, 1973
niger Peucedanum palustre Apiaceae Wagner, 1973
cicadifrons Cistus sp. Cistaceae Wagner, 1973
erythroleptus Calamintha nepeta Lamiaceae Wagner, 1973
leucocephalus Campanula spp. Campanulaceae Göllner-Scheiding, 1972
luridus Calicotome villosa Fabaceae Pericart, 1965
luridus Jasione montana Campanulaceae Göllner-Scheiding, 1970
luridus Jasione montana Campanulaceae Göllner-Scheiding, 1972
Strongylocoris
luridus Jasione montana Campanulaceae Wagner, 1973
oberthuri Jasione montana Campanulaceae Wagner, 1973
obscurus Calicotome villosa Fabaceae Pericart, 1965
obscurus Jasione montana Campanulaceae Wagner, 1973
raimondoi Elaeoselinum asclepium Asteraceae Carapezza, 1991
luridus Jasione montana Campanulaceae Wagner, 1973
eyes rounded and substylate, caudally directed, lat- than body in males, in females shorter than body; AI
erally not touching anterolateral margins of prono- short, swollen in punctulata. Thorax (Figs 3, 11A,
tum. Antennae (Figs 3, 11A–C): antennal insertion in C–E): pronotum rectangular, slightly broader posteri-
line with ventral margin of eye; thin, slightly longer orly, pronotal collar indistinct, callosite region weakly
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 concave, club directed laterally; phallotheca elongate-
rounded, posterior margin weakly concave medially; oval; ductus seminis relatively long and slender, with
mesoscutum not visible; scutellum small and trian- flexible ribbing ending prior to ductus seminis;
gular; metathoracic spiracle large, exposed, with secondary gonopore sclerotized, horseshoe-shaped
evaporative bodies posteriorly and dorsally; MTG basally, opens into less sclerotized, textured hood;
efferent system broad, swollen, triangular, evapora- endosoma with single, large, elongate S-shaped
tive area extending to metathoracic spiracle, ostiole spicule, basally twisted and bifurcate. Female genita-
prominent, directed laterally, peritreme irregularly lia (Fig. 12D, E): sclerotized rings of DLP anteriorly
rounded. Hemelytra (Figs 3, 11A): coleopteroid, shell- narrowed and subcontiguous medially, highly convo-
like; clavus and corium fused without subdivided luted and heavily sclerotized, narrow, posteriorly
appearance; posterior margins convex; elongate with recurved forming sinuate short U-shaped canal,
abdominal tergites VII-IX exposed. Legs (Figs 3, posterior margin of VLP sclerotized; posterior wall
11F): metafemora incrassate; hindlegs elongate. Male of bursa copulatrix simple, weakly sclerotized later-
genitalia (Fig. 12A–C): pygophore conical; both ally, without processes, medially membranous; vesti-
parameres of roughly equal size; left paramere bular region slightly tumescent, bilaterally symme-
L-shaped, weakly concave, apex of apophysis hooked, trical, opening bordered by short, narrow paired
sensory lobe swollen; right paramere clubbed, sclerites.
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 1962: 385 (tribal placement); Wagner, 1973: 107
species and is restricted to eastern Turkey and the (description); Schuh, 1995: 46 (cat.)
Ural Mountains in Russia.
Diagnosis: Chorosomella is distinguished from all
Included species: Barbarosia declavata (Seiden- other halticines by the following combination of char-
stücker, 1962) Turkey acters: gracile body; elongate appendages; rectangu-
Barbarosia punctulata (Reuter, 1901)* Russia: late head; frons with conical projection; vertex with
Ural Mts.; Turkey midline sulcus; bucculae narrow; eyes distant from
anterior margin of pronotum; AI greatly elongate and
Biology and host plant associations: Barbarosia thick; labium short, reaching mesocoxae; MTG exter-
punctulata has been collected from a single species of nal efferent system obsolete; pretarsi with pulvilli;
grass (Elymus junceus) (Kerzhner & Yachevskii, 1967; single elongate endosomal spicule.
Wagner, 1973; Schuh, 1995) (Table 1).
Redescription: Male macropterous, female micropter-
Remarks: The biology of Barbarosia is poorly known. ous. Coloration (Fig. 3): pale yellow-brown. Surface
It is most similar to Euryopicoris, sharing punctate, and vestiture (Figs 3, 13A–D): smooth, almost gla-
coleopteroid, shell-like forewings. Additionally, only in brous. Antennae clothed in very short adpressed
these two genera does the evaporative area of the spinose setae, becoming most dense on AII through
MTG extend upward nearly to the dorsum (24-2). The AIV; femora with very short, adpressed spinose setae
apparent proclivity for grass-feeding by Barbarosia primarily on dorsal surface; tibiae with slightly longer,
and Euryopicoris is also suggestive of a close relation- semi-erect spine-like setae, shorter than leg width.
ship. However, the male aedeagus and female DLP Structure: head (Figs 3, 13A, B): rectangulate, elon-
are unique in Barbarosia, and these characters alone gate, length greater than head height; vertex flat with
are deserving of genus-group status. thin medial longitudinal sulcus, posterior margin not
carinate; frons with conical projection, extending
beyond clypeus; posterior margin of eyes removed from
CHOROSOMELLA HORVÁTH (FIGS 3, 13–14) anterior margin of pronotum; genae height subequal to
Chorosomella Horváth, 1906: 545 (gen. nov.; type eye height; mandibular and maxillary plates small;
species: Chorosomella jakowleffi Horváth, 1906 by bucculae extremely narrow; antennae (Figs 3, 13A, B):
monotypy); Carvalho, 1959: 283 (cat.); Kerzhner, inserted at eye midheight; elongate, slender, longer
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 cylindrical, thin; pretarsi with fleshy pulvilli. Abdomen
combined; AII approximately twice as long as AI; AIII (Fig. 1): elongate in both sexes. Male genitalia
slightly longer than first; AIV short. Labium (Fig. 13A, (Fig. 14A–C): pygophore elongate, posterior margin
B): short, nearly reaching mesocoxae. Thorax (Figs 3, laterally with pair of caudally projecting processes;
13A–D): pronotum narrow, trapezoidal, posteriorly parameres roughly equal in size; left paramere narrow,
only slightly wider than anterior margin, nearly flat, weakly bent, weakly concave, apex of apophysis bifid;
collar absent, callosite region distinct, lateral margins right paramere weakly concave, without apical apo-
rounded, posterior margin medially concave; mesoscu- physis; phallotheca elongate, narrow and simple,
tum elongate; scutellum short, apex blunt, rounded; apically slender; ductus seminis elongate, very thin;
metathoracic spiracle a thin slit, with evaporative secondary gonopore weakly sclerotized, opening
bodies along posterior margin, MTG external efferent into elongate tubular spine-like spicule, reaching
system obsolete. Hemelytra (Fig. 3): males macropter- apex of phallotheca. Female genitalia (Fig. 14D):
ous, costal margin concave, becoming very narrow sclerotized rings thin, elongate, ribbon-like, very
towards apex; membrane with single cell, apex reaches weakly sclerotized. DVP and VLP membranous,
abdominal tergite VII, minor cell obsolete; females without sclerotization; posterior wall of bursa copula-
micropterous, hemelytra translucent; without cuneal trix entirely membranous, without processes or
fracture or membrane; only reaching abdominal modifications vestibular region symmetrical. Vestibu-
tergite III. Legs (Figs 3, 13A, B, E): very long, lar region symmetrical, very weakly swollen, with
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 first moved Chorosomella to the Halticini, whereas
anteriorly. Wagner (1973) regarded the halticines as a subfamily,
and erected a new tribe for Chorosomella: the
Diversity and distribution: Chorosomella is comprised Chorosomellini. Most authors follow Kerzhner (1962),
of two species from central districts of the Palae- who based his decision on various characters he
arctic region, including the Russian Crimea and considered typical of the Halticini: L-shaped left
Kazakhstan. paramere, spoon-shaped right paramere, membra-
nous aedeagus with spicula, and tall genae, and sup-
Included species: Chorosomella horvathi Kiritshenko, ported by their close relationship with graminoids. In
1911 Kazakhstan examining Chorosomella we find that some of these
Chorosomella jakowleffi Horváth, 1906* Russia character evaluations are disputable; compared
to other Halticini the left paramere is much less
Biology and host plant associations: Chorosomella angular; the secondary gonopore is unlike that of any
jakowleffi is known to breed and feed on grasses other Halticini, and the genae are considerably
(Kerzhner & Yachevskii, 1967; Wagner, 1973; Schuh shorter. Additionally, the long, gracile body shape,
& Lattin, 1980) (Table 1). long thin legs, pale coloration, the unique shape of the
head, and the structure of the female genitalia, par-
Remarks: Chorosomella was previously placed in the ticularly the sclerotized rings, are also unlike all
Mirinae: Stenodemini (Horváth, 1906) based on other Halticini.
superficial similarities, such as the elongate body and In our unweighted phylogenetic analysis
appendages, synonymous with many grass-feeding Chorosomella groups with Dimorphocoris and Com-
true bugs. Reuter (1910) transferred the genus to positocoris (Fig. 1) by several homoplasies, including
the Orthotylini, but Carvalho (1952) subsequently an overall nonblack coloration (1-0), a slightly elon-
restored it to the Stenodemini. Kerzhner (1962) gate head (10-1), and the extreme brachyptery of
females, with the wings reduced to wing pads (37-3). beyond the scope of this study, as we only had access
However, under implied weighting this relationship to a few species of the speciose Dimorphocoris.
dissolves, with Chorosomella now appearing as sister
to Plagiotylus in a clade that also includes Strongy-
DAMPIERELLA TATARNIC (FIGS 3, 17–18)
locoris. This clade is supported by the lack of met-
athoracic evaporative area (24-0) and the extreme Dampierella Tatarnic, 2009: 47 (gen. nov.; type
reduction of the metathoracic scent gland itself (23-0). species: Dampierella schwartzi Tatarnic, 2009).
Based on the current data it is unclear which of these
competing topologies more closely reflects the true Diagnosis: Dampierella is superficially similar to
placement of Chorosomella. Halticus but may be distinguished by the much
shorter head height, the presence of a well-
developed endosoma with a single spine-like spicule,
COMPOSITOCORIS SCHWARTZ, SCHUH & TATARNIC and lack of wing reduction in either sex. The geni-
(FIGS 3, 15–16) talia of both sexes are typical of the Halticini, with
the left paramere bearing a thin apical apophysis,
Compositocoris Schwartz, Schuh & Tatarnic, 2008: 23
the right paramere elongate and clubbed, the female
(gen. nov.; type species: Compositocoris senecionus
posterior wall lacking inter-ramal lobes, and the
Schwartz et al., 2008 by original designation)
vestibulum weakly sclerotized and bilaterally sym-
metrical. However, unlike in most other halticines,
Diagnosis: Compositocoris is recognized by the follow- the base of the left paramere is not elongate, a trait
ing combination of characters: strongly sexually shared with another endemic Australian genus,
dimorphic, males macropterous and elongate, females Goodeniaphila.
ovoid and micropterous; bright green coloration;
females with apically stellate bristle-like setae; Diversity and distribution: Dampierella is a mono-
medial margins of first gonapophyses forming vesti- typic genus, with all specimens collected from two
bular opening symmetrical and weakly sclerotized. localities in north-west Western Australia.
This genus is very similar to Dimorphocoris but is
distinguished by the shorter genal height, coloration, Included species: Dampierella schwartzi Tatarnic,
and the apically stellate setae of the females. 2009* Australia
Diversity and distribution: Compositocoris is a mono- Biology and host plant associations: Dampierella has
typic genus, known only from Namaqualand, South only been collected from the plant Dampiera incana
Africa. var. incana R.Br. (Goodeniaceae) (Table 1).
Remarks: This is one of two halticine genera endemic

Included species: Compositocoris senecionus Schwartz to Australia, the other being Goodeniaphila, both of
et al., 2008* South Africa which are found only on the plant family Goodeni-
aceae. Our phylogeny places these genera as sister
Biology and host plant associations: This species is taxa, based on several homoplasious characters and
known from numerous species of asterids belonging to one unambiguous synapomorphy (48-2: the base of
the genera Senecio, Leyesera, and Ursinia (Aster- the right paramere is elongate, whereas that of left
aceae) (Table 1). paramere is much shorter). A complete description
can be found in Tatarnic (2009).
Remarks: Compositocoris is very similar to Dimorpho-

coris in many aspects, including overall male and DASYSCYTUS FIEBER (FIGS 3, 19)
female body form, the presence of paddle-like sclerites Dasyscytus Fieber, 1864: 84 (gen. nov.; type species:
apical to the secondary gonopore, and the patches of Dasyscytus sordidus Fieber, 1864 by monotypy);
honeycombed texturing on the head and thorax (2-1: Kirkaldy, 1906: 131 (cat.); Reuter, 1910: 162 (cat.);
the only unambiguous synapomorphy linking these as Oshanin, 1910: (cat.); Carvalho, 1952: 73 (cat.); Car-
sister taxa in our phylogeny). Only the bright colora- valho, 1955: 68 (key); Carvalho, 1958: 8 (cat.);
tion, shorter head, and stellate bristle-like setae in Wagner, 1973: 30 (descr.); Schuh, 1995: 47 (world
the female distinguish Compositocoris. A more thor- cat.).
ough examination of Dimorphocoris is necessary to Kilicanata Seidenstücker, 1956: 66 (gen. nov., type
identify the range of morphological variation and to species: Kilicanata pilifera = junior synonym of Dasy-
determine if Compositocoris is congeneric; this was scytus: syn. by Wagner, 1960: 99)
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 3.2–3.5 mm; macropterous female 3.3–3.5 mm;
the presence of setae on the hemelytral membrane in coleopteroid female 2.5–3 mm. Coloration (Fig. 3):
macropterous males. mostly dark brown to black, with some orange-brown,
yellow-brown, and yellow markings. Macropterous
Redescription: Males macropterous, females male mostly orange-brown with yellow and tan mark-
coleopteroid or macropterous. Macropterous male ings. Surface and vestiture (Fig. 3): macropterous –
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 angular, occupies lower third of tergite, ostiole lateral,
setae, including on membrane of hemelytron. tongue-shaped, peritreme rounded, directly above
Coleopterous – with fewer setae, surface glossy. ostiole, surrounded by evaporative bodies. Hemelytra
Antennae with white simple setae and several spine- (Fig. 3): macropterous – long and parallel-sided,
like setae, most prominent on AI. Pronotum weakly extending beyond apex of abdomen, cuneus long and
rugulose. Structure: head (Fig. 3): transverse, thin; membrane with two cells. Coleopterous –
approximately as tall as broad; width slightly wider reduced to shell-like pads without division, posterior
than anterior margin of pronotum; vertex trans- margins running diagonally from apex of claval com-
versely sulcate, posterior margin carinate, weakly missure to lateral margin of abdomen, abdominal
wraps around pronotum with eyes touching prono- tergites IV and V partially covered, VI to apex
tum; genae height slightly greater than eye height; exposed. Legs: long and slender; metafemora not
mandibular plate somewhat swollen in macropterous swollen. Abdomen (Fig. 3): elongate and narrow in
male; maxillary plate carinate. Antennae: insertion in macropterous males, not surpassing hemelytral
line with lower margin of eye; AI weakly swollen, margins; rounded in macropterous females, laterally
length approximately equal to eye height; AII weakly extending beyond lateral hemelytral margins.
apically clavate, slightly more than three times AI Coleopterous: rounded pear-shape. Male genitalia
length. Thorax (Fig. 3): pronotum trapezoidal, ante- (Fig. 19A–C): pygophore conical; left paramere api-
rior weakly declivent in lateral view, collar not visible, cally folded, sensory lobe short and narrow, apophysis
callosite region poorly defined, posterior of humeral sinuate and tapering, angled downwards, apically
angles weakly depressed, posterior margin weakly bifid; right paramere spoon-shaped, angled laterally;
concave; metathoracic spiracle large and exposed, phallotheca apically narrow, apicodorsally slightly
narrowly bordered with evaporative bodies running compressed; ductus seminis elongate with flexible
downwards but not confluent with metathoracic ribbing; base of secondary gonopore a sclerotized ring;
evaporative area; MTG external efferent system tri- apically irregularly sclerotized, transversely broad-
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 adjacent portion of DLP weakly upturned; posterior
and apical lobe densely covered with fields of short wall of bursa copulatrix membranous with thin,
teeth. Female genitalia (Fig. 19D–F): DLP transverse, weakly sclerotized rods converging dorsally, ventral
sclerotized rings indistinct, triangular-oval, lateral margin weakly sclerotized; margins of vestibulum
margin most strongly sclerotized, lateral margin and symmetrical and very weakly sclerotized.
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 sion of subfamily placement); Kerzhner & Schuh,
found throughout much of the Mediterranean region, 1995: 8 (revised subfamily placement, disc.); Schuh,
but has not been collected in Syria, Egypt, Italy, and 1995: 47 (world cat.)
southern France (Wagner, 1973).
Diagnosis: Distinguished from all other Halticini by
Included species: Dasyscytus sordidus Fieber, the following combination of characters: small, gracile
1864* Mediterranean body; rounded head with large eyes; semitranslucent
hemelytra; aedeagus with extremely narrow, convo-
Biology and host plant associations: Dasyscytus is luted ductus seminis, endosoma with irregular scle-
found in vegetation at the edges of rivers and ditches, rotized mass apical to secondary gonopore, possibly
with adults collected in April and May (Wagner, formed by multiple strand-like spicules.
1973). No clear host records exist, although specimens
have been collected under Rhanterium epapposum Redescription: Small and fragile; macropterous. Col-
(Asteraceae) in north-east Arabia (Linnavuori, 1986). oration (Fig. 3): glossy; head, pronotum, and abdomen
dark brown, contrasting with mostly pale antennae,
Remarks: The monotypic genus Dasyscytus differs hemelytra, and legs. Hemelytra translucent. Surface
from all other halticines by the presence of setae on and vestiture (Figs 3, 20A–D, G, H): body sparsely
the hemelytral membrane in macropterous individu- clothed in long pale or dark simple setae and regions
als (character 39-1), and the distinct folding of the left of very short, dense, adpressed setae. Head mostly
paramere (Fig. 19B: not coded in this analysis). In our smooth with a few long setae, clypeus and labrum
phylogenetic analysis Dasyscytus is sister to more densely covered with shorter setae; posterior of
Piezocranum, based on the thin hind femora of gena with dense short setae. Antenna clothed in
females (41-0) and the unbroken sclerotization of the short, slender spine-like setae; AI with a few thin
apical region of the ductus seminis up to the second- spines, slightly longer than width of segment. Prono-
ary gonopore (62-1). tum with sparse covering of longer simple setae,
collar with short dense pilosity, thoracic pleura with
dense, short pilosity, Hemelytron sparsely covered
DICYPHOPSIS POPPIUS (FIGS 3, 20–21) with long decumbent setae. Femora clothed in long,
Dicyphopsis Poppius, 1914: 11 (gen. nov.; type species: reclining, simple to weakly spinose setae; tibiae with
Dicyphopsis nigriceps Poppius, 1914 by monotypy); semi-erect spine-like setae. Structure: head (Figs 3,
Carvalho, 1952: 72 (cat.); Carvalho, 1955: 58 (key); 20A–D): rounded, weakly transverse, posterior
Carvalho, 1958: 191 (cat.); Cassis, 1986: 164 (discus- margin of vertex weakly convex; wider than tall,
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 angular; metathoracic spiracle small, with narrow
approximately equal to eye height; eyes rounded, band of evaporative bodies above; MTG efferent
sessile, not touching anterior of pronotum; frons system narrow and angled posteriorly, ostiole a ver-
broadly rounded, bulging. Labium (Fig. 20C, D): tical slit, peritreme tongue-shaped, running along
extends to metacoxae. Antennae (Figs 3, 20A–D): ventral margin of tergite, bordered above by narrow
insertion in front of ventral margin of eye; long and band of evaporative bodies. Hemelytra (Fig. 3): semi-
thin, AI slightly longer than eye height. Thorax translucent, lateral margins parallel, cuneus long and
(Figs 3, 20D, E): pronotum trapezoidal, steep, collar narrow, membrane extends beyond abdomen. Legs
broad and flat, callosite region faintly medially (Figs 3, 20A, B, F): long and slender; metafemur
depressed, humeral angles rounded, posteriorly slightly incrassate; pretarsi without fleshy pulvilli.
deeply depressed, lateral and posterior margins Abdomen: narrow, parallel-sided, in males slightly
concave; mesoscutum visible; scutellum thin and tri- broadened at pygophore. Male genitalia (Figs 20G, H,
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, broad, irregular sclerotized mass near secondary
sensory lobe broad, apophysis short, tapering, with gonopore, possibly made of many long, thin sclerites.
small subapical keel, apically hooked; right paramere Female genitalia (Fig. 21D–F): DLP membranous,
longer than left, straight, spoon-shaped; phallotheca sclerotized rings transverse, medially subcontiguous,
elongate-oval, simple; ductus seminis elongate and subrectangulate, very weakly sclerotized; VLP simple,
constricted, with flexible ribbing; secondary gonopore with medial margin sclerotized and dentate; posterior
ill defined, weakly sclerotized; endosoma large, with wall of bursa copulatrix weakly sclerotized anteriorly,
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 Remarks: The monotypic genus Dicyphopsis was
bilateral swellings; margins of vestibulum symmetri- described from a single female specimen of Dic. nig-
cal with very weak sclerotization. riceps collected in Tanzania (Poppius, 1914). Unfortu-
nately the holotype was destroyed during shipment to
G. Cassis (Cassis, 1986; Kerzhner & Schuh, 1995),
Diversity and distribution: Dicyphopsis includes only
leaving no existing specimens of this species. Kerzh-
one described species from Tanzania (Kilimanjaro),
ner & Schuh (1995) identified a number of specimens
though undescribed specimens matching the original
that they considered congeneric (although not conspe-
description and considered congeneric have been col-
cific) collected in Zaire by N. A. Weber in 1948, and
lected in Zaire and Ghana (Kerzhner & Schuh, 1995).
Ghana by R. T. Schuh and J. A. Slater in 1967. We
have only seen the Ghana specimens, and although
Included species: Dicyphopsis nigriceps Poppius, the habitus is not typical of the Halticini, we agree
1914 Kilimanjaro. with their assertion that the genitalia is of the Hal-
ticini type, albeit with some minor differences. Most Diagnosis: Recognized by the following combination
notably, in the aedeagus the ductus seminis is of characters: female always brachypterous, male
extremely narrow, and the spicule form is unique. In macropterous or brachypterous, macropterous indi-
the female however, the margins of the vestibulum viduals elongate and parallel-sided; body brown with
are symmetrical, a character consistent throughout yellow-brown and tan markings; frons, pronotum,
the tribe. It is worth noting that what we call Dicy- and thoracic pleura with faint, symmetrical honey-
phopsis superficially appears identical to what Lin- combed texturing; eyes round and bulging, substy-
navuori (1994) identifies as the Orthotylini genus late to stylate; posterior margin of vertex with
Nycticapsus Poppius (D. Forero, pers. comm.). Both row of coarse black setae; endosoma with sclerites
genera were described in Poppius (1914), with neither apparently originating from apex of secondary
one illustrated. As we have not seen specimens of gonopore.
Nycticapsus, for the moment we do not know whether
Poppius described the same species twice, whether Redescription: Dimorphic, males macropterous or
Linnavuori misidentified specimens of Dicyphopsis, or brachypterous, females brachypterous. Coloration
whether these are in fact distinct genera. As the (Fig. 3): Highly variable, but generally a combination
genitalia are clearly of the Halticini rather than the of dark brown to black with yellow, yellow-brown, or
Orthotylini type, we retain Kerzhner & Schuh’s tan markings. Surface and vestiture (Figs 3, 22A–E,
(1995) initial placement of these specimens. Phyloge- G, H): body mostly smooth and impunctate, with
netic analysis places Dicyphopsis in a clade of prima- some bilaterally symmetrical, honeycomb-like textur-
rily African taxa, defined by the presence of paired ing on vertex, frons, pronotum and thoracic pleura,
bilateral pillow-like swellings on the posterior wall these patches sometimes congruent with contrasting
(87-1). These taxa also share similar metathoracic colour patterning. Most of body densely covered with
scent gland structure, with the peritreme finger-like, short, pilose setae, much less densely covered with
angled back and resting along the caudal margin of longer simple and spine-like setae; sometimes also
the metepimeron (25-3), a condition shared only by with long, silvery scale-like setae; posterior of vertex
Piezocranum and Coridromius (could not be deter- with a transverse row of coarse black setae; antennae
mined in Namaquacapsus because of condition of and tibiae with short, semi-reclining bristle-like
specimen). setae, AI typically with several longer, thicker
bristles. Structure: head (Figs 3, 22A–D): transverse,
DIMORPHOCORIS FIEBER (FIGS 3, 22–23) wider than tall; gena height equal or greater than eye
Dimorphocoris Reuter, 1890: 253 [gen. nov.; type height; frons mostly flat and sloping, sometimes with
species: Orthlocephalus signatus Fieber, 1861 subse- shallow medial sulcus continuing onto vertex; vertex
quent designation (junior synonym of Phytocoris gra- sometimes with bilateral depressions adjacent to
cilis Rambur, 1839)]; Reuter, 1891: 83, 159 (gen. nov., eyes, posterior margin of vertex straight, not
key); Hueber, 1906: 5, 6 (key, descr.); Kirkaldy, 1906: upturned or carinate, weakly concave to weakly
131 (cat.); Oshanin, 1910: 784 (cat.); Reuter, 1910: 147 convex; eyes round and bulging, substylate to stylate,
(cat.); Poppius, 1914: 85 (key, descr.); Bergevin, 1916: projecting laterally beyond and not touching antero-
61 (dimorphism); Stichel, 1933: 235 (key); Hedicke, lateral margins of pronotum, sometimes angled
1935: 59 (key); Kiritshenko, 1951: 126 (key); Wagner, upwards. Antennae (Figs 3, 22B, C): insertion in line
1952: 96, 107 (key, descr.); Carvalho, 1952: 73 (cat.); with or below ventral margin of eye; long, thin, and
Wagner, 1954: 221 (key to spp.); Carvalho, 1955: 67 cylindrical, always longer in males but generally not
(key); Carvalho, 1958: 8 (cat.); Lindberg, 1956: 56 longer than body length; AI sometimes swollen,
(disc., key); Wagner, 1961: 50 (diag., key); Kerzhner, always longer than eye height; AIV shorter than AIII.
1964b: 964 (diag., key); Wagner & Weber, 1964: 276 Thorax (Figs 3, 22A, D, E): pronotum campanulate in
(syn., descr., key); Kerzhner, 1964a: 121 (key to spp. macropterous males, rectangular in brachypterous
from USSR); Wagner, 1965a: 135 (disc., descr.); individuals, gently sloping to nearly level, collar thin,
Wagner, 1965b: 57 (illustrated key); Wagner, 1969: 79 often obscured by strongly tumescent callosite region,
(modified key); Dimorphocoris (Pedetocoris) Kerzhner, posterior margin straight to concave; mesoscutum
1970: 634 (subgen. nov., diag., disc., key); Wagner, visible in macropterous males, steeply posteriorly
1973: 57 (descr., key); Linnavuori, 1992: 216 (key to declivent; scutellum often medially tumescent in mac-
lateralis group); Ehanno, 1994a: 10–37 (illustrated ropterous males, broad and flat in brachypterous indi-
disc. of morphology, incl. MG, FG); Schuh, 1995: 48 viduals; metathoracic spiracle prominent, diagonal,
(world cat.). with evaporative bodies; MTG external efferent
Homoeocoris Fuente, 1918: 236 (gen. nov. type system tear-shaped, occupying posterior half of
species: Orthocephalus tristis Fieber, 1861: syn. by metepisternum, ostiole narrow and orientated vent-
Wagner, 1955: 241); Carvalho, 1958: 8 (cat.) rolaterally, peritreme elongate, weakly depressed,
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. Abdomen (Fig. 3): elongate in macropterous males,
Hemelytra (Fig. 3): Macropterous male: elongate, elongate-oval to pear-shaped in brachypterous indi-
parallel-sided, cuneus long and thin; membrane with viduals. Male genitalia (Figs 23A–C, 22G, H): pygo-
two cells, extends beyond tip of abdomen. Brachypter- phore triangular in lateral view; left paramere
ous: usually reduced to an undivided pad, sometimes L-shaped, sensory lobe rounded and weakly swollen,
with faint clavus. Legs (Figs 3, 22F): long, particu- apophysis tapering, apex often hooked or weakly
larly in macropterous males; pretarsi without pulvilli. bifid; apex of right paramere angled and spoon- or
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 Included species: Dimorphocoris abutilon Wagner,
apophysis; aedeagus fully enclosed within phalloth- 1966 France
eca; ductus seminis elongate, with flexible ribbing; Dimorphocoris albipilis Kerzhner, 1964
secondary gonopore sometimes cup-shaped, often Kazakhstan
weakly sclerotized and ill-defined, sometimes with Dimorphocoris alpinus Poppius, 1910 Kenya
scale-like texturing; endosoma with either complex, Dimorphocoris argaeicus Hoberlandt, 1956
serrate, sclerotized outpocketings (lateralis group) or Turkey
with single or paired sclerotized, pedunculate pro- Dimorphocoris asanovae Kerzhner, 1964 Palae-
cesses, often serrate, emanating from dorsal wall of arctic region (western)
secondary gonopore (Dim. gracilis group). Female Dimorphocoris atrans Kerzhner, 1970 Russia
genitalia (Fig. 23D, E): sclerotized rings small, widely Dimorphocoris beieri Wagner, 1965 the former
separated, mostly round to elongate-oval, lateral Yugoslavia
margins generally not curved upwards but lateral Dimorphocoris bleusei Puton, 1898
margin of adjacent portion of dorsal labiate plate Mediterranean
typically upturned; posterior wall of bursa copulatrix Dimorphocoris carayoni Ehanno & Ribes, 1994
variable, often divided into two bilaterally symmetri- France
cal regions of weak sclerotization, one anterior and Dimorphocoris cilix Seidenstücker, 1962 Turkey
one posterior, sometimes partly covered with fields of Dimorphocoris concii Tamanini, 1972 Italy
minute spines; vestibular opening symmetrical, with Dimorphocoris constantini Ehanno, 1994a France
medial margins of first gonapophyses weakly swollen Dimorphocoris debilis (Reuter, 1880)
and weakly sclerotized. Mediterranean
Dimorphocoris distylus Seidenstücker, 1964
Turkey
Diversity and distribution: With 58 species this is the Dimorphocoris dupuisi Ehanno, 1993a France
largest genus of Halticini. Dimorphocoris is found Dimorphocoris durfortae Ehanno & Ribes, 1994
predominantly in the Mediterranean region but is Spain
also known to extend into North and East Africa, Dimorphocoris eckerleini Wagner, 1965
Central Asia, and Sakhalin Island in the North Mediterranean
Pacific (Ehanno, 1991, 1993, 1994a, b; Wagner, 1973; Dimorphocoris ehannoi Ribes & Ribes’ 2001
Linnavuori, 1992; Ehanno & Ribes, 1994). The genus Morocco
is subdivided into two groups, the Dim. gracilis and Dimorphocoris fuscus Joakimov, 1909* Palaearc-
Dim. lateralis groups. tic region (western)
Dimorphocoris gallicus Wagner, 1965 France Poaceae (Table 1). Some species have also been col-
Dimorphocoris goulae Ehanno & Ribes, lected from higher level angiosperms, with Dim. osel-
1994 Spain lai collected from unidentified species of Lotus and
Dimorphocoris gracilis (Rambur, 1839)* Algeria; Trifolium (Fabaceae) (Tamanini, 1976), Dim. tristis
Spain found on Anthemis sp. (Asteraceae) (Linnavuori,
Dimorphocoris josephinae Ehanno & Ribes, 1992), and Dim. seidenstueckeri collected from uni-
1994 Spain dentified species of Asteraceae, Teurcrium (Lami-
Dimorphocoris lateralis Reuter, 1901 Palaearctic aceae), and Trifolium (Fabaceae) (Linnavuori, 1984)
region (western) (Table 1).
Dimorphocoris lividipennis Reuter, 1903 Spain Many species, especially of the Dim. gracilis group,
Dimorphocoris longiceps Wagner, 1968 Morocco have restricted mountainous distributions, whereas
Dimorphocoris lurensis Wagner, 1957 France others are more widely distributed in littoral meadows
Dimorphocoris marci Rizzotti Vlach, 1998 Italy and on the grassy steppes of North Africa (Wagner &
Dimorphocoris marginellus (Puton, 1887) Algeria; Weber, 1964; Ehanno & Ribes, 1994). It is thought the
Morocco restricted ranges of many species may be the result of
Dimorphocoris mariae Linnavuori, 1952 the high degree of brachyptery expressed in the genus,
Mediterranean particularly in females (Linnavuori, 1992).
Dimorphocoris matocqui Ehanno, 1993 France
Dimorphocoris mongolicus Kerzhner, 1970 Remarks: Dimorphocoris is divided into two species
Mongolia groups: the Dim. lateralis group in which both sexes
Dimorphocoris mutatus Seidenstücker, 1964 Italy are brachypterous, and the Dim. gracilis group in
Dimorphocoris obachi Ehanno & Ribes, 1994 Spain which males are macropterous whereas females are
Dimorphocoris osellai Tamanini, 1976 Italy brachypterous (Linnavuori, 1992). Linnavuori (1992)
Dimorphocoris pedetes Kerzhner, 1964 further subdivided the former into three ‘phylogenetic
Kazakhstan subgroups’ (although no phylogenetic results were
Dimorphocoris pericarti Tamanini, 1972 France; presented). The highly speciose Dim. gracilis group is
Italy; Spain also comprised of several distinct subgroupings,
Dimorphocoris poggi Carapezza, 2002 Italy which are dealt with in detail by Ehanno (1993,
Dimorphocoris puigmalis Tamanini, 1976 France 1994a, b) and Ehanno & Ribes (1994).
Dimorphocoris punctiger (Horváth, 1881) Israel; Members of the Dim. gracilis group are most
Syria similar in appearance and in genitalic form to the
Dimorphocoris putoni (Reuter, 1882) Andorra; newly described monotypic genus Compositocoris
France Schwartz et al., 2008: see appendix). In both cases the
Dimorphocoris pygmaeus Wagner, 1955 France males are macropterous whereas the females are
Dimorphocoris remanei Wagner, 1965 Spain brachypterous, and the genitalia of both sexes are
Dimorphocoris robustus Wagner, 1957 France very similar. In particular, the presence of two paddle-
Dimorphocoris ruffoi Tamanini, 1971 Italy shaped sclerotized processes in the endosoma sug-
Dimorphocoris sari Linnavuori, 1992 Greece gests a close relationship. Only the presence of
Dimorphocoris satyriscus (Scott, 1870) Spain apically stellate bristles on the body of the female and
Dimorphocoris saulii Wagner, 1965 Italy; Slov- the structure of the posterior wall set Compositocoris
enia; the former Yugoslavia apart. Although there is strong evidence to synony-
Dimorphocoris schmidti (Fieber, 1858) Ukraine mize Compositocoris with Dimorphocoris, we refrain
Dimorphocoris seidenstueckeri Linnavuori, 1984 from doing so without examination of more specimens
Iraq of Dimorphocoris.
Dimorphocoris servadeii Tamanini, 1982 Italy Although we dissected few specimens of Dimorpho-
Dimorphocoris tamaninii Ehanno, 1993 France coris, Ehanno & Ribes (1994; Ehanno, 1994a, b)
Dimorphocoris tauricus (Horváth, 1880)* Russia provide detailed illustrations of many species, includ-
Dimorphocoris tiberghieni Ehanno, 1993 France ing both male and female genital structure, which
Dimorphocoris tomasii Tamanini, 1971 Italy greatly assisted in informing this redescription.
Dimorphocoris toros Seidenstücker, 1962 Turkey
Dimorphocoris tristis (Fieber, 1861) Spain
Dimorphocoris tuatayae Wagner, 1965 Morocco ECTMETOPTERUS REUTER (FIGS 3, 24–25)
Ectmetopterus Reuter, 1906: 59 [gen. nov. type
Biology and host plant associations: Most species are species: Ectmetopterus angusticeps Reuter, 1906 by
thought to feed on grasses (Wagner, 1973) with the monotypy (junior synonym of Halticus micantulus
majority of host records coming from the family Horváth, 1905)]; Oshanin, 1910: 802 (cat.); Reuter,
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, Redescription: Macropterous. Coloration (Fig. 3): gen-
1952: 73 (cat.); Carvalho, 1955: 65 (key); Carvalho, erally black or brown, sometimes with brown and
1958: (10 cat.); Schuh, 1995: 52 (world cat.). orange-brown markings. Surface and vestiture
(Figs 3, 24A–H): body smooth and impunctate, with
Diagnosis: Similar to Halticus and Microtechnites, but both simple reclining setae and short scale-like setae.
readily distinguished by its trifurcate left paramere Setae on legs and antennae semi-erect and spine-like.
and broad and flat right paramere. Structure: head (Figs 3, 24A–D): transverse, slightly
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 athoracic spiracle tear-drop shaped, surrounded with
shallow transverse depression, posterior margin thin thin border of evaporative bodies extending up along
and carinate; genae height greater than eye height; posterior margin of mesepimeron; MTG external
eye short and tall; frons steeply sloping; clypeus efferent system tear-drop shaped, angled posterodor-
slightly projecting; maxillary plate swollen; buccula sally, ostiole rounded and orientated laterally, peri-
thin. Labium (Fig. 24C, D): LI swollen and elongate, treme tongue-like, tapered at apex. Hemelytra
approximately as long as genae height; labium (Fig. 3): costal margins evenly curved, broader sub-
reaches metacoxae. Antennae (Figs 3, 24A–D): inser- apically; clavus apically broadened; R + M vein long,
tion in line with lower half of eye; longer than body, extending almost to cuneus; membrane surpasses
cylindrical; AI approximately as long as eye height; apex of abdomen. Legs (Figs 3, 24F): metatibiae
AII almost 1.5¥ as long as pronotal width; antennae slightly swollen; pretarsi with pulvilli. Abdomen:
longer than body length. Thorax (Figs 3, 24A, D, E): parallel-sided, slightly broader in females. Male geni-
pronotum trapezoidal, short, steep, collar thin, lateral talia (Figs 25A–C, 24G, H): pygophore conical, poste-
margins weakly concave, humeral angles rounded, rior margin sinuate, deeply concave below left
posterior margin thin, weakly declivent, straight to paramere; parameres with long, thick base; left
weakly medially cleft; mesoscutum not visible; met- paramere broad, L-shaped and trifurcate, with apex
of first branching apophysis hooked, sometimes with typy); Stichel, 1933: 235 (key); Hedicke, 1935: 58
apophyses slightly twisted; right paramere straight, (key); Kiritshenko, 1951: 127 (key); Wagner, 1952: 96,
flat, and broad, extends out of pygophore, apicolateral 105 (key, descr.); Carvalho, 1955: 66 (key); Euryopi-
margins sometimes strongly recurved; phallotheca coris Reuter, 1875b: 24 (typogr. error, key); Reuter,
elongate-oval, tapering to apex; ductus seminis trun- 1875b(1): 87; (2): 99 (key, descr.); Reuter, 1891: 60, 158
cate, with flexible ribbing; secondary gonopore basally (descr., key); Hueber, 1906: 4, 11 (key, descr.);
sclerotized, apically indistinct; endosoma with weakly Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 790 (cat.);
sclerotized medial structure extending apically from Reuter, 1910: 147 (cat.); Carvalho, 1952: 73 (cat.);
secondary gonopore. Female genitalia (Fig. 25D, E): Carvalho, 1958: 11 (cat.); Wagner & Weber, 1964: 274
DLP unmodified; sclerotized rings widely separated, (descr.); Wagner, 1973: 51 (descr.); Schuh, 1995: 53
elongate-ovoid, diagonal, tapered laterally and medi- (world cat.).
ally, lateral margins slightly upturned; DLP with a
thin transverse sclerotized band adjacent to juncture Diagnosis: Distinguished from other genera by the
with posterior wall; margins of VLP sclerotized, medi- combination of punctate, coleopteroid hemelytra and
ally membranous; posterior wall of bursa copulatrix curved pro- and mesotibiae.
membranous, medially and laterally with linear fields
of spines, laterally with paired, swollen tumescences. Redescription: Both sexes usually coleopterous, males
Vestibulum symmetrical, swollen, and uniformly rarely macropterous. 3–4.5 mm. Coloration (Fig. 3):
weakly sclerotized. nearly uniform glossy black, sometimes with yellow-
brown at apices of femora and antennal segments,
Diversity and distribution: All six species of Ectme- hemelytra of macropterous individuals reddish-
topterus are found in eastern Asia. brown; eyes red. Surface and vestiture (Figs 3, 26A–F,
H): surface glossy; head smooth with radiating folds
Included species: Ectmetopterus bicoloratus Kulik, emanating from centre of vertex; pronotum irregu-
1965* comb. nov. East Russia larly rugulose anteriorly, punctate posteriorly; scutel-
Ectmetopterus comitans Josifov & Kerzhner, 1972* lum rugulose; hemelytron punctate. Body with sparse
comb. nov. East Russia distribution of simple setae, mostly on ventral part of
Ectmetopterus fuscosus Zou, 1985 comb. head and ventral and lateral surfaces of body; anten-
nov. China nae and legs with semi-erect spine-like setae inter-
Ectmetopterus maculipes Zou, 1985 comb. spersed with longer, thicker spines, particularly on AI
nov. China and tibiae. Structure: head (Figs 3, 26A–E): trans-
Ectmetopterus micantulus (Horváth, 1905)* East verse, broader than anterior of pronotum; vertex flat,
Asia posterior margin thin, weakly bisinuate; eyes small
Ectmetopterus niger Zou, 1985 comb. nov. China and rounded, substylate; head height approximately
three times eye height; frons broadly rounded, con-
Biology and host plant associations: No information fluent with clypeus; buccula thin; labium (Fig. 26D,
available. E): reaches mesocoxae, LI short and thick. Antennae
(Figs 3, 26A–E): insertion in front and below ventral
Remarks: Based on similarities in male genitalia,
margin of eye; short, approximately as long as body;
herein we transfer to Ectmetopterus five species of
AI swollen, about two times length of eye height; AIII
Halticus: Ec. bicoloratus, Ec. comitans, Ec. fuscous,
shorter than AIV. Thorax (Figs 3, 26A, B, E–G):
Ec. maculipes, and Ec. niger (all comb. nov.). As with
pronotum short, trapezoidal and nearly flat, posterior
Ec. micantulus, all of these species have East Asian
slightly broader than head, collar absent, callosite
distributions and very similar overall structure. The
region weakly tumescent, lateral margins rounded,
trifurcate left paramere, long and flat right paramere,
humeral angles broadly rounded, posterior margin
and structure of the aedeagus clearly place them in
weakly medially cleft; mesoscutum not visible; scutel-
this genus. This redescription is based on examina-
lum transverse; metathoracic spiracle prominent and
tion of Ec. micantulus, Ec. comitans, and Ec. bicolora-
elongate, surrounded by a thin band of evaporative
tus. Although we have not seen the three Chinese
bodies; MTG external efferent system broad, triangu-
species of Zou (1985), based on the descriptions and
lar, and swollen, ostiole vertically orientated, opens
illustrations of the parameres they are congeneric
laterally, peritreme tear-shaped, orientated vertically,
with the above.
surrounded by evaporative bodies. Hemelytra (Figs 3,
26A): macropterous: extending beyond apex of
EURYOPICORIS REUTER (FIGS 3, 26–27) abdomen; costal margin weakly convex, nearly
Euryopicoris Reuter, 1875b: 11 (gen. nov.; type straight; membrane with single cell. Coleopterous
species: Capsus nitidus Meyer-Dür, 1843 (by mono- individuals: long, partially covering tergite VI; undi-
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 pygophore conical, genital opening narrow, posterior
weakly rounded. Legs (Figs 3, 26H): metafemora margin weakly sinuate, with shallow concavity below
slightly incrassate; fore- and mesotibiae distinctly left paramere; both parameres long and thin basally;
curved and thickened; pretarsi with fleshy pulvilli. left paramere with tumescent sensory lobe, apophysis
Abdomen (Fig. 3): pear-shaped in both sexes, becom- angled upwards, apically hooked; right paramere
ing conical at apex. Male genitalia (Fig. 27A–C): with angled apical club, apicolateral margin recurved;
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 Diversity and distribution: There are two species
seminis elongate, basally with flexible ribbing, of Euryopicoris, with Euryopicoris fennicus known
subapically with elongate, weakly sclerotized only from Finland and Euryopicoris nitidus broadly
section lacking flexible ribbing; secondary gonopore distributed across central Europe into Siberia.
rounded, sclerotized, and U-shaped with prised
operculum and fine scale-like texturing; endosoma Included species: Eu. fennicus Wagner, 1954 Finland
with a single spine-like spicule. Female genitalia Eu. nitidus (Meyer-Dür, 1843)* Central and North
(Fig. 27E, F): sclerotized rings widely separated, Europe; Siberia
strongly sclerotized, oval, lateral, and medial margins
and adjacent portions of DLP strongly upturned; Biology and host plant associations: Euryopicoris
lateral margins of DLP moderately sclerotized, medi- nitidus is found in mountain meadows and is said
ally with transverse bilaterally sulcate sclerotized to live on grasses. Adults are found from June to August.
plate (similar to the inter-ramal bridge in Labops), This species overwinters as eggs (Wagner, 1973). There
middle of plate a small carinate point projecting is no biological information for Eu. fennicus.
into bursa copulatrix; posterior margin of VLP
weakly sclerotized; posterior wall of bursa copu- Remarks: Euryopicoris is similar to Anapus and Bar-
latrix simple and plate-like, medially membranous, barosia in general structure. Although the secondary
laterally weakly sclerotized; vestibulum symme- gonopore of Euryopicoris is most similar to that of
trical, opening bordered by short, narrow paired Anapus, the lack of prominent sclerotized processes on
sclerites. the female posterior wall, the punctate hemelytra, and
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 phyletic clade, which together form the sister group to
apart. Euryopicoris shares punctate coleopteroid hem- Myrmecophyes, Labops, and Scirtetellus.
elytra and a bulging frons with Barbarosia, but the
structure of the DLP in Euryopicoris is relatively simple GOODENIAPHILA TATARNIC (FIGS 1, 28–29)
in contrast to the complexity expressed in Barbarosia. Goodeniaphila Tatarnic, 2009: 52 (gen. nov.; type
Results of both unweighted and implied weighting phy- species: Goodeniaphila cassisi Tatarnic, 2009: 53 by
logenetic analyses placed these three genera in a mono- original designation).
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; Goodeniaphila schuhi Tatarnic, 2009* Australia
vestibulum weakly sclerotized and mostly symmetri-
cal, except for small sclerotized knob on left margin.
Goodeniaphila most closely resembles Strongylocoris, Biology and host plant associations: Goodeniaphila
but can be distinguished by the presence of a well- has been collected exclusively on the plant family
developed MTG external efferent system (absent in Goodeniaceae.
Strongylocoris). The dull, olive coloration of the hem-
elytra and the male and female genitalia set it apart
from all other Halticini. Remarks: Along with Dampierella, Goodeniaphila is
endemic to Australia. Both weighted and unweighted
Diversity and distribution: This genus is known from phylogenies place these two in a well-supported
two species collected in Western Australia, the North- sister-taxon relationship. The male genitalia of Good-
ern Territory, and South Australia. eniaphila is mostly typical of the Halticini – the left
paramere bears an apically hooked apophysis, the
Included species: Goodeniaphila cassisi Tatarnic, right paramere is elongate and clubbed, the ductus
2009* Australia seminis is ribbed, the secondary gonopore bears scale-
like texturing, and the endosoma is well developed (Western Hemisphere key); Linnavuori, 1994: 5
and contains several sclerotized spicules distal to the (African key); Cassis & Gross, 1995: 186 (synonymy;
secondary gonopore. However, the base of the left Australian cat.); Schuh, 1995: 53 (synonymy; world
paramere is short (48-2: shared with Dampierella) cat.).
rather than elongate as in other halticines. The Eurycephala Laporte, 1832: 36 (gen. nov.; Lygaeus
female genitalia is also mostly typical of the tribe – luteicollis Panzer, 1805 by subsequent designation
well-developed sclerotized rings with lateral margins Kirkaldy, 1906: 170); Brullé, 1836: 140 (descr.);
and adjacent portion of DLP weakly upturned, poste- Spinola, 1840: 191 (descr.); Blanchard, 1840: 140
rior wall without inter-ramal lobes – however, the (descr.); Reuter, 1891: 17 (synonymy); Carvalho, 1958:
sclerites of first gonapophyses are nearly symmetrical 12 (cat.)
(as in other halticines) but for a minute sclerotized Halticocoris Douglas & Scott, 1865: 478
lobe projecting from the left side (Fig. 29G). This lobe (unnecessary nom. nov. for Halticus Hahn nec
is not seen in other halticines but is present in some Haltica Illiger [Coleoptera]); Carvalho, 1958: 13
Orthotylini, albeit in a much more pronounced state (cat.);
(Schaffner & Schwartz, 2008: 41, fig. 19c). For a com- Halticiellus Villiers, 1952: 186 (gen. nov.; type
plete description of Goodeniaphila see Tatarnic species: Halticus tibialis Reuter, 1891 by monotypy);
(2009). Carvalho, 1958: 12 (synonymy)
Diagnosis: Halticus is characterized by a combination

HALTICUS HAHN (FIGS 3, 30–32) of the following: length of first labial segment less
Astemma Latreille, 1829: 199 (nom. nud.); junior than gena height; elongate, flattened right paramere;
homonym of Astemma Lepeletier & Serville, 1825 left paramere with twisted apophysis; scoop-shaped
[Heteroptera: Pyrrhocoridae]; type species: Acanthia secondary gonopore and vestigial endosoma lacking
pallicornis Fabricius, 1794 = Cicada aptera Linnaeus, spicules; large, thin sclerotized rings.
1758 by monotypy; Westwood, 1840: 121 (description);
Amyot & Serville, 1843: 284 (description) Redescription: Usually small and robust, macropter-
Halticus Hahn, 1833: 113 (gen. nov.; type species: ous or coleopteroid. Coloration (Fig. 3): almost all
Acanthia pallicornis Fabricius, 1794 = Cicada aptera species black, sometimes with paler yellow, orange,
Linnaeus, 1758 by monotypy); Burmeister, 1835: 277 red, or brown colouring on head, antennae, and legs,
(descr.; Spinola, 1850: 43 (key); Fieber, 1858: 312 rarely uniformly brown. Surface and vestiture (Figs 3,
(key); Fieber, 1861: 69, 281 (key; descr.); Reuter, 30A–E, G, H): variable, generally glossy, sometimes
1875a: 23 (key); Reuter, 1875b: 86, 89 (key; descr.); wholly impunctate, in others pronotum is shallowly
Saunders, 1875: 287 (key); Atkinson, 1890: 117 (cat.); punctate; frons and vertex with faint creases radiat-
Reuter, 1891: 17, 157 (descr.; key); Saunders, 1892: ing from centreline. Body generally with two forms of
267 (key; descr.); Distant, 1904: 479 (descr.); Kirkaldy, setae – thin and simple and white, flat, and scale-like;
1906: 130 (cat.); Hueber, 1906: 2, 42 (key; descr.); antennae and legs with short semi-erect spines and
Oshanin, 1910: 802 (cat.); Reuter, 1910: 147 (cat.); spine-like setae. Structure: head (Figs 3, 30A, B, D):
Poppius, 1914: 84 (key; descr.); Van Duzee, 1916: 211, transverse and short, ranging from broader than tall
374 (cat.); Knight, 1923: 498 (key); Blatchley, 1926: to taller than broad; vertex with transverse sulcus,
800 (eastern US species key); Stichel, 1933: 235 posterior margin carinate, concave, generally broader
(Palaearctic key); Hedicke, 1935 (Palaearctic key); than anterior of pronotum; head height variable,
Knight, 1941: 17, 75, 77 (Illinois key; descr.); Hsiao, usually > two times eye height; eyes sometimes wrap-
1942: 253 (Chinese key); China, 1943: 268 (nomencla- ping around pronotum, sometimes touching; eyes tall
ture); Zimmerman, 1948: 199 (Hawaiian key); Slater, and short; clypeus narrow, dorsal margin weakly
1950: 45 (female genitalia); Kiritshenko, 1951: 127 sulcate; mandibular plate depressed; maxillary plate
(USSR key); Wagner, 1952: 95, 96 (Palaearctic key; raised and carinate. Labium (Fig. 30D): length vari-
descr.); Carvalho & Leston, 1952: 245 (British key); able, LI greatly swollen. Antennae (Figs 3, 30A–D):
Carvalho, 1952: 74 (list); Carvalho, 1955: 65 (key); insertion typically in line with or slightly below lower
Carvalho, 1956: 51 (key Micronesian spp.); Carvalho, margin of eye; long, thin, and whip-like; AI slightly
1958: 12 (cat.); Southwood & Leston, 1959: 244 thicker than other segments, about as long as eye
(British key); Kelton, 1959: 33, Fig. 81 (male genita- height. Thorax (Figs 3, 30A, D–F): pronotum trapezoi-
lia); Wagner, 1961: 48 (diag.; Palaearctic key); Kerzh- dal (macropterous) to subrectangular (coleopterous),
ner, 1964a: 968 (diag.; USSR key); Maldonado, 1969: weakly declivent, laterally sloping, collar thin and flat
57 (diag.; US key); Alayo, 1974: 15 (diag.); Schuh, to absent; callosite region ill-defined to obsolete, pos-
1974: 29 (discussion); Wagner, 1973: 6 (descr.; Palae- terior margin thinly carinate, straight to weakly
arctic key); Kelton, 1980: 194 (diag.); Henry, 1983: 610 medially cleft; metathoracic spiracle prominent, nar-
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 exter- side steeply sloping; corium laterally sloping; mem-
nal efferent system swollen, occupying lower half of brane and cuneus deflected. Coleopterous: broadly
metepimeron, ostiole opens laterally, peritreme rounded, laterally and posteriorly sloping; claval
rounded, orientated posteriorly, surrounded by evapo- suture and medial fracture faint, cuneal fracture very
rative bodies. Hemelytra (Fig. 3): macropterous: short. Legs: long, metafemur greatly swollen; pretarsi
costal margin weakly curved, broadest above cuneal with fleshy pulvilli. Male genitalia (Figs 30H, 31A–E):
fracture; clavus longitudinally folded, with lateral pygophore conical, posterior margin sinuate, with
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; phalloth- sometimes angled laterally at midpoint, apex rounded
eca simple and elongate-oval; left paramere apically with a small apicolateral apophysis; ductus seminis
twisted with sinuate apophysis; right paramere attenuate, with flexible ribbing; secondary gonopore
longer than left, extends out of pygophore, broad, flat, partly sclerotized, in form of elongate scoop, nearly
filling phallotheca; endosoma extremely reduced, Remarks: Most recent definitions of Halticus are
without spicules. Female genitalia (Figs 31F–H, 32A– broad, inclusive of great morphological heterogeneity
C): dorsal labiate plate mostly membranous; sclero- (e.g. Wagner, 1973), and pay little regard to genitalic
tized rings large, transverse, medially subcontiguous, features. This is the first study to account for the
sclerotized margins thin, lateral margins weakly morphological diversity exhibited across species of
upturned; VLP weakly sclerotized on posterior Halticus, and to examine this in a phylogenetic
margin, laterally sometimes with paired, scimitar- context. As a consequence we have subdivided Halticus
shaped sclerotizations, curving laterally and reaching sensu lato into three genera. Based primarily on male
up to dorsal labiate plate; posterior wall of bursa and female genitalic characters, and supported by our
copulatrix variable, for the most part membranous, phylogeny, we remove ten species examined that do not
often with narrow paired sclerotized bands or rods follow the model of the type species, H. apterus (left
convergent anteriorly; vestibulum symmetrical, with paramere L-shaped with apically curved apophysis;
medial margins of first gonapophyses swollen out- ductus seminis short; secondary gonopore elongate and
wards and sclerotized, vestibular opening rounded scoop-shaped, endosoma obsolete and without spi-
and strongly sclerotized. cules; sclerotized rings large and subcontiguous).
H. bractatus, H. canus, H. chrysolepis, H. minutus,
and H. spegazzini are removed, and along with both
Diversity and distribution: Halticus has 16 species species of Cafayatina, form the resurrected genus
and a mainly Holarctic distribution, with most species Microtechnites. Similarly, the Asian species H. bicol-
confined to the Palaearctic. A single species, Halticus oratus, H. comitans, H. maculipes, H. niger, and
tibialis, is found in Africa, South-East Asia, and the H. fuscous are all transferred to Ectmetopterus.
south-west Pacific. We have not seen all species of Halticus, and it is
therefore possible that with further examination
Included species: Halticus apterus (Linnaeus, other species may yet be removed from the genus.
1758)* Palaearctic
Halticus asperulus Horváth, 1898 Armenia LABOPS BURMEISTER (FIGS 4, 33–36)
Halticus beganus Linnavuori, 1984 Iraq
Labops Burmeister, 1835: 279 [gen. nov.; type species:
Halticus darbandikhaus Linnavuori, 1984 Iraq
Labops diopsis Burmeister, 1835 by monotypy (junior
Halticus henschii Reuter, 1888 Italy
synonym of Capsus sahlbergii Fallén, 1829)]; Fieber,
Halticus insularis Usinger, 1946 Mariana Is.;
1858: 316 (key); Fieber, 1860a: 294; 1861 (key, descr.);
Guam; Korea
Flor, 1860: 448 (key); Thompson, 1871: 432 (key);
Halticus intermedius Uhler, 1904 USA
Walker, 1873: 44 (cat.); Reuter, 1875a: 24 (key);
Halticus luteicollis (Panzer, 1804)* Europe;
Reuter, 1875b(1): 86 (2):97 (key, descr.); Provancher,
Mediterranean
1887: 135 (descr.); Atkinson, 1890: 120 (cat.); Reuter,
Halticus macrocephalus Fieber, 1858
1891: 80, 160 (descr., key); Hueber, 1906: 5 (key);
Mediterranean
Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 786 (cat.);
Halticus major Wagner, 1951 Europe
Reuter, 1910: 148 (cat.); Van Duzee, 1916: 211 (key);
Halticus obscurior Kerzhner & Muminov, 1974
Van Duzee, 1917: 373 (cat.); Knight, 1922: 258 (key, n.
Asiatic Russia
spp.); Blatchley, 1926: 797 (key, spp. nov.); Stichel,
Halticus puncticollis Fieber, 1870 Montenegro
1933: 235 (key); Knight, 1941: 81; Slater, 1950: 51
Halticus pusillus (Herrich-Schäeffer, 1835)*
(genitalia); Kiritshenko, 1951: 126 (key); Carvalho,
Palaearctic
1952: 74 (cat.); Slater, 1954: 57 (note); Carvalho, 1955:
Halticus rugosus Reuter, 1894 Syria
66 (key); Carvalho, 1958: 18 (cat.); Kerzhner, 1964a:
Halticus saltator (Geoffroy, 1785) Europe
964 (diag., key); Kelton, 1980: 189 (diag., key); Kerzh-
Halticus tibialis Reuter, 1891* Africa; South-East
ner, 1988: 43 (illustrated key to east Asian spp.);
Asia, South Pacific
Schuh, 1995: 58 (world cat.).
Ophthalmocoris Zetterstedt, 1838: 280 gen. nov.
Biology and host plant associations: Limited host (syn. by Herrich-Schäeffer, 1850:166).
records exist for Halticus. Halticus apterus and H. be-
ganus have both been collected from the family Diagnosis: Readily distinguished from other Halticini
Fabaceae (Vicia striata and Phaseolus vulgaris, by the following combination of characters: tall head
respectively). Halticus beganus has also been col- and stylate eyes; hemelytra of brachypterous indi-
lected from rotten oak (Fagaceae) (Henry & Smith, viduals nearly covering abdomen; aedeagus with
1979). Halticus pusillus is said to live on and beneath mass of thin, needle-like spicules suspended in the
Galium verum (Rubiaceae) (Wagner, 1973) (Table 1). membrane anterior to secondary gonopore; posterior
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 scler- larly on vertex, along midline, and on genae, mandibu-
ites similar to those found in other Orthotylinae; DLP lar and maxillary plates. Hemelytra black, sometimes
often with heavily sclerotized inter-ramal bridge. with white or yellow along costal margins. Surface and
vestiture (Figs 4, 33A–H): clothed in simple setae and
Redescription: Coloration (Fig. 4): mostly dark, dull pale, scale-like setae. Head smooth and setaceous.
brown to black, usually with yellow, orange, or reddish Antennae and legs with erect and semi-erect spines,
markings. Head: black with yellow markings, particu- particularly on AI and tibiae. Posterior of pronotum
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.
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 rounded; membrane with two cells. Legs: long, metafe-
setae or both simple and scale-like setae. Structure: mur not greatly swollen; pretarsi without fleshy pul-
body elongate-oval. Head (Figs 4, 33A–C): transverse villi. Abdomen: elongate-oval. Male genitalia
and tall; genae height > two times eye height; eyes (Figs 33F–H, 34A–E, 36A–D): pygophore simple and
stylate, projecting laterally and slightly upward, conical, ventral margin extending slightly caudad, left
extending well beyond the anterolateral margins of the ventrolateral margin sulcate below left paramere; left
pronotum; frons steep, with faint folds radiating from paramere either straight or with obtusely angled apo-
midline; mandibular plate sometimes tumescent, espe- physis, apophysis weakly curved to strongly angled
cially males. Labium (Fig. 33C): variable, sometimes apically, sensory lobe often prominent; right paramere
reaching metacoxae; LI swollen. Antennae (Figs 4, extends caudally out of pygophore, considerably larger
33A–C): insertion below eye; shorter than insect; AI than left paramere, with long base, apex with laterally
swollen, shorter to longer than pronotum, approxi- deflected concave club; apex of phallotheca with
mately twice as long as eye height. Thorax (Figs 4, serrate, plate-like process of variable size projecting
33A, C, D): pronotum campanulate, collar thin and flat, from the left of phallothecal opening; ductus seminis
callosite region well defined, posterior margin weakly elongate, secondary gonopore sclerotized, vaguely
concave; mesoscutum broad; scutellum small, triangu- bowl-shaped with a prised operculum, with scale-like
lar; metathoracic spiracle small and exposed, thinly texturing along opening margin; endosoma with tight
surrounded with evaporative bodies; MTG external bundle of sclerotized needles, sometimes also with
efferent system broad, swollen and triangular, encom- other sclerotized processes, including thin strips of
passing lower half of tergite, ostiole prominent and sclerotized teeth. Female genitalia (Figs 34D–G,
elongate, peritreme tongue-shaped, situated above 35A–B, 36E–G): sclerotized rings large, elongate-
ostiole in middle of evaporative area, angled rearward elliptical, diagonal, medially subcontiguous, lateral
parallel to posterior margin of tergite. Hemelytra margins somewhat upturned; margin of VLP adjoining
(Fig. 4): both sexes with brachypterous and macropter- rami sclerotized, sometimes medially with paired
ous morphs, degree of brachyptery varying across teeth, lateral-most region of VLP joined with rami to
species (Slater, 1954); length variable in brachypterous form paired, medially projecting, sclerotized processes,
morphs but generally covering much if not all of the sometimes covered with fields of spines; DLP some-
abdomen, cuneus sometimes differentiated, membrane times with heavily sclerotized, bilaterally ventrally
either absent or vestigial; in macropterous morphs pointed inter-ramal bridge, sometimes reduced to
hemelytra extend beyond abdomen, margins gently paired pincer-like lateral sclerites, a simple band of
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 Labops nivchorum Kerzhner, 1988 Russia
bursa copulatrix with bilaterally symmetrical inter- Labops sahlbergii (Fallén, 1829)* Scandinavia;
ramal lobes of varying complexity projecting from Siberia (Ural)
posterior wall; margins of first gonapophyses sym- Labops setosus Reuter, 1891 Russia
metrical, swollen, and sclerotized. Labops tumidifrons Knight, 1922* Canada
Labops utahensis Slater, 1954* USA
Diversity and distribution: Labops is comprised of 12 Labops verae Knight, 1929 USA
species and exhibits a Holarctic distribution.
Biology and host plant associations: Species of Labops
Included species: Labops bami Kulik, 1979 Russia are associated with grasses in arid conditions (Slater,
Labops brooksi Slater, 1954 Canada 1954). In North America, L. hesperius has been
Labops burmeisteri Stål, 1858 Holarctic recorded on Agropyron cristatum, various range
Labops chelifer Slater, 1954 Canada grasses, Rosa arkansana (Kelton, 1980), Koeleria cris-
Labops hesperius Uhler, 1872* Canada; USA tata, Poa secunda, Stipa comata, Stipa williamsi,
Labops hirtus Knight, 1922 Canada; USA Hordeum sp., Triticum sativum (Mills, 1939: in Slater,
1954), and wheat (Mills, 1941), whereas L. hirtus has similar to Halticus in salient features, but is
been recorded on wheat in Montana (Mills, 1939). In separated from the latter by the elaborate endo-
Yakutia, Vinokurov (1979) recorded the genus on somal spicules and the widely separated sclerotized
cereals and sedges (Carex stenophylla, Critesion rings.
jubatum, and Leymus sp.), and suggested that it may
decrease productivity of pastures and hay fields Redescription: Male and female macropterous and
(Table 1). brachypterous morphs, sometimes males macropter-
ous only (e.g. M. bractatus), very small to small,
Remarks: Labops has previously been treated as its 1.5–3 mm. Coloration (Fig. 4): body mostly dark
own tribe, Labopini, owing to various characteristics brown with yellow-brown and yellow markings.
unique to the genus (Knight, 1923). Slater was the Surface and vestiture (Figs 4, 37A–H): body sparsely
first to note the unusual female genitalia of Labops – clothed in semi-erect simple setae intermixed with
both the posterior wall (1950, 1954) and the inter- decumbent, scale-like setae; legs with semi-erect
ramal bridge (1954). Slater (1950) questioned spines. Head mostly smooth, posterior of pronotum
whether the sclerotized inter-ramal tumescences on weakly rugulose. Structure: head (Figs 4, 37A–C):
the posterior wall might be homologous to the inter- transverse, elongate in lateral view; posterior
ramal lobes (i.e. K-structures) found in other Ortho- margin of vertex sublinear, carinate; head slightly
tylinae – the absence of which has been considered a broader than anterior margin of pronotum; frons
synapomorphy for the Halticini (Schuh, 1974). convex; height of genae approximately equal to
However, with the discovery of inter-ramal tumes- height of eye; eyes sessile; maxillary plates some-
cences in both Anapus and Scirtetellus, we consider times carinate; bucculae narrow; antennae (Figs 4,
these to be homologous in origin. Despite the presence 37A–C): antennal insertion in front of eye; cylindri-
of inter-ramal lobes in Labops, Scirtetellus, and cal, variable length; AI subequal to eye height,
Anapus, our phylogeny instead places Labops as weakly swollen; AII-AIV thinner. Labium (Fig. 37C):
sister to Myrmecophyes based on four unambiguous reaching metacoxae; LI slightly enlarged; thorax
characters, including one synapomorphy (56–1: apex (Figs 4, 37A, C–E): pronotum trapezoidal, collar
of phallotheca with a prominent flange). present and thin, callosite region weakly delineated,
The inter-ramal bridge (Kullenberg, 1947; Slater, posterior margin convex, humeral angles rounded,
1950, 1954) is another interesting structure otherwise in lateral view pronotum of macropterous individu-
absent in the Halticini (although Anapus, Barbarosia, als gently sloping, nearly vertical in brachypterous
and Euryopicoris possess a sclerotized plate on the individuals; mesoscutum not visible in both mac-
DLP, which may be homologous). ropterous and brachypterous individuals; scutellum
flat, apically rounded; metathoracic spiracle small
uniformly bounded by evaporative bodies, sometimes
MICROTECHNITES BERG (FIGS 4, 37–38) extending to mesepimeral knob; MTG external effer-
Microtechnites Berg, 1883: 73 [gen. nov.; type species ent system broad, pear-shaped, small, confined to
Capsus pygmaeus Berg, 1879 (junior homonym of medial third of metespisternum, peritreme oval,
Capsus (Deraeocoris) pygmaeus Berg, 1878) = Halti- medially positioned. Hemelytra (Fig. 4): macropter-
cus spegazzinii Berg, 1883 by monotypy]; Reuter, ous: lateral margins weakly arcuate; embolium wid-
1910: 149 (cat.); Carvalho, 1952: 74 (synonymy with ening towards cuneal fracture; clavus broad, claval
Halticus Hahn); Carvalho, 1958: 13 (cat.); Cassis & commissure elongate; median flexion line half length
Gross, 1995: 186 (cat.; synonymy); Schuh, 1995: 53 of claval suture; cuneus transverse; membrane with
(cat.; synonymy). Reinstated status. one cell, minor cell obsolete, membrane extending
Cafayatina Carvalho & Carpintero, 1986: 611 (gen. beyond apex of abdomen. Brachypterous: rounded
nov.; type species: Cafayatina altigena Carvalho & and shell-like, undivided, posterior margins weakly
Carpintero, 1986 by original designation); Schuh, rounded, wholly or partly covering abdominal tergite
1995: 46 (cat.). syn. nov. VI. Legs (Figs 4, 37F): metafemora elongate, incras-
sate; metatibiae elongate; pretarsi with fleshy pul-
Diagnosis: Microtechnites is recognized by the follow- villi. Abdomen: short, lateral margins parallel,
ing combination of characters: macropterous and slightly broader in female. Male genitalia (Figs 37G,
brachypterous morphs; dark ground colour; moderate H, 38A–C): pygophore short, conical, genital aper-
distribution of white scale-like setae; eyes sessile; ture large, posterior margin symmetrical, concave;
parameres subequal in size; left paramere L-shaped; left paramere L-shaped, sensory lobe small, apophy-
male endosoma with numerous elongate spicules, sis broad, apically arcuate; right paramere subequal
often serrate; and, female sclerotized rings widely to left paramere length, spoon shaped; phallotheca
separated, diagonally orientated. Microtechnites is simple; ductus seminis short; secondary gonopore
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 sclerotized, entire, without medial flange; posterior
spicules, most spicules elongate, sometimes serrate. wall of bursa copulatrix subtriangular, weakly scle-
Female genitalia (Fig. 38D–F): sclerotized rings rotized, anteriorly with paired short digitiform pro-
moderately sclerotized, widely separated, lateral cesses; vestibulum symmetrical and weakly
margins with adjacent portion of DLP weakly sclerotized, anteriorly with paired sclerites converg-
upturned; margin of VLP adjacent to rami narrowly ing posteriorly, forming a Y-shape.
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 Included species: Microtechnites altigena (Carvalho &
to the Western Hemisphere, with most species found in Carpintero, 1986)* comb. nov. Argentina
Central and South America. It is the only halticine Microtechnites bractatus (Say, 1832)* Neotropical
genus found in the Neotropical region, including its and Nearctic regions
distribution in the Caribbean and the Galapagos = Rhinacloa citri Ashmead, 1887: 155 (syn. Reuter,
Islands (Henry & Wilson, 2004). Only Mi. bractatus is 1909)
known in the Nearctic region, being widespread in = Halticus minutus Uhler, 1890 nec Reuter, 1884
eastern Canada and the USA and extending as far (junior primary homonym of Halticus minutus
south as Argentina. Microtechnites chrysolepis is Reuter, 1884)
endemic to Hawaii and its distributional isolation is = Halticus uhleri Giard, 1892:81 (new name for
significant. We do not regard this disjunction as nec- Halticus minutus Uhler – syn. Parshley, 1915)
essarily anomalous, as the derivation of Hawaii’s Microtechnites canus Distant, 1893* (restored status,
fauna from the Western Hemisphere has been postu- Henry, 1983: 607) Guatemala; Mexico; Nicaragua
lated for the orthotyline genus Sarona (Asquith, 1995). Microtechnites chrysolepis (Kirkaldy, 1904)* Hawaii
Microtechnites inesalti (Carvalho & Carpintero, Diplacus Stål, 1858: 183 [gen. nov. (n. preocc. by the
1992)* comb. nov. Argentina nomen nudum Diplacus Rafinesque, 1815 (Ascidiae)];
Microtechnites spegazzinii Berg, 1883* Neotropi- – (syn. by Reuter, 1890: 252); Reuter, 1875a: 87 (key);
cal region 2:100 (descr.); Reuter, 1875b: 25 (key); Atkinson,
1890: 117 (cat.); Carvalho, 1958: 139 (cat.)
Biology and host plant associations: Host plants have
Diagnosis: Easily recognized by the following charac-
been recorded only for the ubiquitous Mi. bractatus;
ters: strongly myrmecomorphic, with extreme sta-
Maldonado (1969) recorded two cucurbit species as
phylinoidy in females and most males; tall genae in
host plants, whereas Linnavuori (1984) recorded
both sexes; hemelytra in macropterous individuals
Mi. bractatus on sweet potato (Ipomoea batatas)
without cuneus, membrane without cells.
(Table 1). This species, which is commonly known as
the garden fleahopper, has been recorded as a pest
Redescription: Coloration (Fig. 4): mostly black, some-
from a broad range of crops (Henry, 1983; Carpinera,
times with yellow, red, or orange in parts, sometimes
2001), including solanaceous crops (aubergine,
with cream or yellow along the posterior margin of
tomato, potato, pepper, tobacco), commercial cucurbits
staphylinoid hemelytra, or with mostly pale mac-
(cucumber, pumpkin, squash) and legumes (alfalfa,
ropterous hemelytra. Surface and vestiture (Figs 4,
beans, clover, cowpeas, peas), as well as beets,
39A–H): surface mostly glossy. Body clothed in simple
cabbage, celery, lettuce, barley, corn, oats, wheat, and
setae and white, decumbent scale-like setae. Head
cotton. Carpinera (2001) also recorded this species
mostly smooth with a few simple setae; frons with
from a great number of weed species, regarded as
faint rugulose whorls radiating from medial sulcus.
secondary hosts associated with garden vegetables
Antennae with reclining simple setae, AI with several
(Table 1). Microtechnites bractatus is also thought to
erect black spines, usually more in males. Pronotum
be zoophytophagous (Carpinera, 2001). It has five
mostly smooth with a few erect simple setae, usually
instars and overwinters as eggs. Eggs are laid in
basally and apically rugulose; thoracic pleura clothed
stems and leaf petioles.
in very short, fine setae; scutellum weakly rugulose.
Femora with some reclining setae and several black
Remarks: We have removed Microtechnites from syn- spines dorsally; tibiae with erect spines approxi-
onymy with Halticus and reinstated it to generic mately as long as tibial width, anterior tibiae in
status. In addition, Cafayatina is established as a males of some species highly spinose. Structure: both
junior synonym of Microtechnites. Microtechnites is sexes strongly ant-mimetic, hemelytra almost always
distinguished by its unique male genitalia. In Halti- reduced to minute pads, rarely macropterous. Head
cus the aedeagus ends with an elongate scoop-shaped (Figs 4, 39A–C): transverse, wider than pronotum,
secondary gonopore and the endosoma is vestigial taller than wide, genae height > two times eye height;
without spicules, whereas in Microtechnites the sec- eyes round, substylate, not touching pronotum; pos-
ondary gonopore opens into a membranous endosoma, terior margin of vertex weakly concave, not carinate;
which contains several complex spicules and sclero- frons nearly flat, steeply declivent to base of clypeus;
tized serrations. Additionally, the sclerotized rings of clypeus vertical; buccula small and thin. Labium
Microtechnites are generally smaller and more widely (Fig. 39C): surpasses metacoxae; LI swollen. Anten-
separated, in comparison to Halticus where they are nae (Figs 4, 39A–C): insertion well below ventral
transverse, large, and medially subcontiguous. 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,
MYRMECOPHYES FIEBER (FIGS 4, 39–40) 39A, C–E): pronotum quadrate, more campanulate in
Myrmecophyes Fieber, 1870: 253 [gen. nov.; type macropterous males, collar rarely present, callosite
species: Myrmecophyes oschannini Fieber, 1870 by region sometimes slightly swollen, lateral margins
monotypy (junior synonym of Diplacus alboornatus rounded, posterior margin straight or slightly
Stål, 1858)]; Reuter, 1891: 106 (descr.); Kirkaldy, concave; mesoscutum greatly swollen; scutellum tri-
1906: 132 (cat.); Oshanin, 1910: 781 (cat.); Reuter, angular, in staphylinoid morphs anterolateral
1910: 148 (cat.); Horváth, 1926: 187 (monograph); margins down-curved, apically blunt; metathoracic
Horváth, 1927: 196 (subgen. nov.); Stichel, 1933: 234 spiracle prominent, encircled with thin ring of evapo-
(key); Kiritshenko, 1951: 126 (key); Carvalho, 1952: rative bodies; MTG external efferent system broad
82 (cat.); Wagner, 1955: 240 (transferred to Halticini); and triangular, evaporative area reaching almost as
Carvalho, 1955: 79 (key); Carvalho, 1958: 139 (cat.); high as metathoracic spiracle, ostiole elongate, verti-
Bykov, 1971: 871 (diag., key); Wagner, 1973: 4 (descr.); cally orientated and opening laterally, peritreme
Schuh, 1995: 59 (world cat.). broad, tear-shaped, and vertical. Hemelytra (Fig. 4):
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 pos- cuneus absent, membrane without closed cells. Legs
terior margin of abdominal tergite II, posterior (Figs 4, 39F): long; protibiae of males in some species
margins rounded, sometimes slightly upturned, some- expanded (e.g. My. macrotractus); metafemur only
times somewhat angular, occasionally claval suture somewhat enlarged; pretarsi without fleshy pulvilli.
forming a raised carina. Macropterous – clavus Abdomen (Fig. 4): cylindrical, pear-shaped or globu-
present, medial fracture and R + M vein obsolete, lar, usually noticeably rounder in females, basal seg-
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 ing from left of apex; ductus seminis elongate, with
narrow waist, terminal segments strongly sloping long subapical region more heavily sclerotized and
posteroventrally; laterotergites upturned. Male geni- lacking flexible ribbing; secondary gonopore sclero-
talia (Figs 39G, H, 40A–C): pygophore broad, strongly tized and rounded, with partly open operculum, opens
tapered; left paramere weakly bent, sensory lobe ventrally, surface with scale-like texture; endosoma
often prominent, apophysis straight or curved, apex with variable number of sclerotized spicules, gener-
generally hooked or bifid; right paramere larger than ally two or three, apically often with one or two
left, sometimes extending caudally from pygophore, whorl-like sclerites armed with fields of spines.
apex deflected laterally and spoon-shaped with or Female genitalia (Fig. 40C–E): sclerotized rings vari-
without a small apical apophysis; phallotheca with able in size and shape, ovoid to elongate-elliptical,
serrate, sclerotized flange (sometimes minute) extend- widely separated, lateral margins and adjacent
portion of DLP upturned; DLP and VLP weakly scle- Myrmecophyes piceus Bykov, 1970 Central Asia
rotized and closely adpressed; posterior wall simple Myrmecophyes tibialis Reuter, 1901 Central Asia
and sheet-like, sometimes bilaterally sclerotized; ves- Myrmecophyes trispiculus Drapolyuk & Kerzhner,
tibulum symmetrical, with medial margins bordering 2000 Kazakhstan
opening sinuous. Myrmecophyes variabilis Drapolyuk, 1989 Azer-
baijan
Diversity and distribution: Myrmecophyes contains 29
species and has a Holarctic distribution, with a single Biology and host plant associations: Myrmecophyes is
species (My. oregonensis) restricted to the western found almost exclusively on grasses. In the USA,
USA and the remainder of species found in the Palae- My. orgenonensis has been collected from Festuca ida-
arctic region. hoensis and Agropyron spicatum (Schuh & Lattin,
1980); Kerzhner & Yachevskii (1967) recorded My. al-
Included species: Myrmecophyes acutifrons Bykov, boornatus on Agropyron spp. in Eastern (former)
1971 Central Asia USSR; Bykov (1971) recorded My. macrotrichus on
Myrmecophyes alacer Horváth, 1926 Festuca and Carex spp.; and My. frontosus has been
Transcaucasia collected from Psathyrostachys juncea (Poaceae)
Myrmecophyes alboornatus (Stål, 1858)* (Drapolyuk & Kerzhner, 2000). A single species has
Palaearctic been collected from the family Asteraceae;
Myrmecophyes armeniacus Drapolyuk, 1989 My. trispiculus is known from the asterid Artemisia
Armenia; Turkey terrae-albae (Drapolyuk & Kerzhner, 2000) (Table 1).
Myrmecophyes dubius Bykov, 1971 Central Asia
Myrmecophyes ermaki Bykov, 1969 Central Asia Remarks: Myrmecophyes is arguably the most con-
Myrmecophyes frontosus Drapolyuk & Kerzhner, vincing ant mimetic genus of Halticini. Based on its
2000 Kazakhstan mymecomorphy, Carvalho (1952, 1955, 1958) errone-
Myrmecophyes gallicus Wagner, 1976 France; ously placed the genus in the Pilophorini. Many char-
Spain acters contradict this assessment (e.g. tall genae,
Myrmecophyes geniculatus Reuter, 1894 Central structure of the male and female genitalia) – and in
Asia any case several other Halticini are also ant mimics –
Myrmecophyes heterocerus Horváth, 1926* leading Wagner (1955) and Schuh (1974) to transfer
Armenia the genus to the Halticini.
Myrmecophyes hirsutiventris Bykov, 1971 Central Schuh & Lattin (1980) provided a brief summary of
Asia the genus. The most recent works on Myrmecophyes
Myrmecophyes kiritshenkoi Horváth, 1926 Central have been Drapolyuk’s (1989) description of three
Asia new species and key to the species of the Caucasus,
Myrmecophyes korschinskii Reuter, 1903 Central and Drapolyuk & Kerzhner’s (2000) description of two
Asia new species from Kazakhstan. The most comprehen-
Myrmecophyes lacteipennis Bykov, 1971 Central sive work remains Bykov (1971), which as Schuh &
Asia Lattin (1980) note, omitted nine previously described
Myrmecophyes latus Wagner, 1975 Yugoslavia species from its key and lacks a formal diagnosis.
Myrmecophyes limbatus (Reuter, 1879) Central Thus, the most recent diagnosis of Myrmecophyes is
Asia that of Wagner (1973), which has been found by us
Myrmecophyes lipskii Bykov, 1971 Central Asia and others (e.g. Schuh & Lattin, 1980) to be both
Myrmecophyes macrotrichus Horváth, 1926 inadequate and inaccurate. Contrary to Wagner, not
Central Asia all species have white markings on the hemelytra, not
Myrmecophyes montenegrinus Wagner, all species exhibit a macropterous form, the first two
1976 Montenegro abdominal segments are not constricted to form a
Myrmecophyes monticola Horváth, 1926 Central waist – in fact, only occasionally is the second
Asia abdominal segment constricted, whereas in many
Myrmecophyes muminovi (Kerzhner, 1964) species the abdomen remains relatively broad at its
Central Asia base – and the opening of the pygophore is not
Myrmecophyes nasutus Drapolyuk, 1989 Georgia directed downwards.
Myrmecophyes nigripes (Reuter, 1879) Central The structure of the ductus seminis and secondary
Asia gonopore of Myrmecophyes are most similar to
Myrmecophyes nitens Bykov, 1971 Central Asia Anapus and Euryopicoris. In particular, in all three
Myrmecophyes oregonensis Schuh & Lattin, genera the distal portion of the ductus seminis lacks
1980* USA flexible sclerotized ribbing. This similarity is reflected
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 Diagnosis: Recognized by the following characters: male
well as Labops and Scirtetellus) identified in our macropterous (female unknown); robust and elongate;
phylogenetic results. It should be noted that although predominantly chestnut-coloured, with corium red at
the two species of Myrmecophyes included in our base and apex; impunctate; body clothed in long, erect
phylogenetic analysis do not form a monophyletic black setae; tibiae without spines; genae height subequal
group, we consider this to be an artefact of our small to eye height; AI slightly swollen, AII apically clavate;
taxon sample for the genus as well as reflective of the pronotum broad; collar flat; pretarsi with minute pulvilli;
limits of our character dataset. left paramere L-shaped, with narrow, prominent sensory
lobe and long, thin, apically hooked apophysis; right
paramere apically spoon-shaped.
NAMAQUACAPSUS SCHUH (FIGS 4, 41–42)
Namaquacapsus Schuh, 1974: 26 (gen. nov.; type Redescription: Coloration (Fig. 4): black with rust-
species: Namaquacapsus melanostethoides Schuh, coloured hemelytra. Surface and vestiture (Figs 4,
1974 by monotypy); Schuh, 1995: 62 (cat.) 41A, B): head and thorax smooth and glossy, remain-
der dull, hemelytra weakly pitted; body clothed with 1974). Based on our phylogeny, Namaquacapsus
long, black, erect setae; setae on antennae and legs belongs to a clade of mostly African taxa that includes
thin and spinose; antennae with short semi-erect Dicyphopsis, Ectmetopterus, Acratheus, and Nan-
setae interspersed with a few long, erect setae; niella, which are united by several characters, includ-
metatibiae without spines. Structure: male mac- ing one synapomorphy: the presence of pillow-like
ropterous, female unknown. Head (Figs 4, 41): swellings on the posterior wall (87-1). As only the
deflexed, short and transverse, broader than anterior male of Nam. melanostethoides is known, the states of
of pronotum; posterior margin of vertex raised and this and other female-specific characters are
carinate; frons broadly convex, steeply declivent; gena unknown for this species. Additionally, because the
height slightly greater than eye height; eyes tall and legs of the sole specimen we examined obscure our
short, substylate, not touching anterior margins of view of the metathoracic scent gland structure, it is
pronotum; buccula narrow. Labium (Fig. 41A): labium uncertain whether or not the peritreme is directed
reaches anterior of metacoxa, LI short and swollen. caudally as a thin strip along the posterior margin of
Antennae (Figs 4, 41A): insertion close to and in line the metepimeron (25-2), as is the case in the other
with lower margin of eye; thin and cylindrical, shorter clade members.
than body length; AI slightly swollen, shorter than
eye height; AII narrow basally, weakly apically
NANNIELLA REUTER (FIGS 4, 43–44)
clavate; AIII and AIV thin. Thorax (Fig. 4): pronotum
trapezoidal, collar flattened, callosite region obsolete, Nanniella Reuter, 1904: 5 (gen. nov.; type species:
anterior of humeral angles depressed, posterior Nanniella chalybea Reuter, 1904 by monotypy);
margin declivent, weakly rounded, thin and carinate; Kirkaldy, 1906: 131 (cat.); Reuter, 1910: 148 (cat.);
mesoscutum concealed by pronotum; scutellum Carvalho, 1952: 76 (syn.); Carvalho, 1958: 61 (cat.);
strongly convex. Hemelytra (Figs 4, 41A): long; costal Schuh, 1974: 28 (diag., disc. of tribal placement);
margin rounded; clavus steeply declivent laterally; Linnavuori, 1975: 48 (disc. of tribal placement, key);
embolium laterally sloping; cuneal fracture deep, Linnavuori, 1994: 9 (descr., key); Schuh, 1995: 62
cuneus long, laterally rounded; membrane with two (world cat.).
cells. Legs (Fig. 41A, B): metafemora only weakly
swollen; pretarsi with minute pulvilli. Male genitalia Diagnosis: Nanniella is distinguished from other Hal-
(Fig. 42A–C): parameres elongate, right longer than ticini by the following combination of characters:
left; left paramere with narrow, projecting sensory metallic black coloration; uniformly coarsely punc-
lobe, arm concave, apically bifid; right paramere tate; left paramere with twisted, somewhat serrate
longer than left, apical club broad, deflected from apophysis; right paramere apicolaterally recurved
midline, concave, with apex weakly swollen: phalloth- with finger-like apical apophysis. Nanniella is most
eca apically constricted, without lateral projections; similar to Acratheus, but can be separated by the
ductus seminis long and narrow, with flexible ribbing, presence of deep punctures on the frons and vertex
subapical portion without ribbing, secondary gonop- and the shape of the parameres.
ore sclerotized, curved and dorsoventrally constricted
at apex, ventral surface with serrate/scale-like Redescription: Length: 2.5–4 mm. Colouration
texture; endosoma with weakly sclerotized folds ema- (Fig. 4): mostly black with yellow-brown or yellow
nating from secondary gonopore, apically with mod- markings. Surface and vestiture (Figs 4, 43A–G):
erately sclerotized serrate and weakly spinose flange. body with metallic lustre; densely covered with small,
Female unknown. slit-like punctures, becoming less dense on abdomen,
venter impunctate; frons with deep punctures. Body
Diversity and distribution: Known only from Nama- covered with long, greyish, semidecumbent, simple
qualand, South Africa. setae; legs with semi-erect, short spine-like setae.
Structure: both sexes macropterous, body elongate
Included species: Namaquacapsus melanostethoides and slightly ovate. Head (Figs 4, 43A–D): transverse,
Schuh, 1974* South Africa slightly broader than tall, broader than anterior of
pronotum, height approximately twice eye height;
Biology and host plant associations: There are no host vertex with shallow transverse sulcus, posterior
or ecological data. margin slightly upturned and weakly carinate,
straight with eyes not touching pronotum; frons
Remarks: Namaquacapsus is known from only two almost flat, steeply declivent; clypeus vertical, not
male specimens. The original description lacked illus- projecting forward; buccula small and narrow. Anten-
trations of the aedeagus, with the endosoma said to nae (Figs 4, 43A–B, D): insertion in front and slightly
be membranous, presumably lacking spicules (Schuh, below midpoint of eye, nearly touching eye; long and
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 metathoracic spiracle elongate and exposed, without
than eye height. Thorax (Figs 4, 43A, B, D–F): prono- evaporative bodies; MTG efferent system narrow,
tum trapezoidal, elongate and bulging, collar broad angled caudally, ostiole nearly ventral, peritreme a
and flat, callosite region poorly defined, lateral thin strip along posteroventral margin of tergite, bor-
margins rounded, posterior of humeral angles weakly dered dorsally by evaporative bodies. Hemelytra
depressed, posterior margin wider than head, medi- (Fig. 4): elongate, lateral margins subparallel, mem-
ally concave; mesoscutum not visible; scutellum flat; brane extends beyond abdomen. Legs (Fig. 4): long
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 sclerotized channel within endosoma. Female genita-
fleshy pulvilli. Abdomen (Fig. 43H): parallel-sided, lia (Fig. 44F–G): DVP and VLP membranous; sclero-
slightly broader at anterior of genital capsule. Male tized rings small, thin, and weakly sclerotized,
genitalia (Figs 43G, H, 44A–E): pygophore basally weakly upturned at lateral margins; posterior wall of
broad, short and conical; both parameres long, sub- bursa copulatrix membranous, covered with dense
equal in length; left paramere elongate, sensory lobe field of minute setae or spines, with very faint lateral
rounded, apical apophysis angled upwards, twisted, swellings; margins of vestibulum weakly swollen,
dorsally dentate, apically bifid; right paramere pro- symmetrical, with four small sclerotized patches.
jecting slightly out of pygophore, broad, laterally
recurved, with curved finger-like apical apophysis;
Diversity and distribution: All six species of Nanniella
phallotheca simple and elongate; ductus seminis and
share an Afrotropical distribution.
secondary gonopore broad and dorsoventrally com-
pressed; ductus seminis attenuate, weakly sclero-
tized, without flexible ribbing; secondary gonopore Included species: Nanniella alkithoe Linnavuori,
indistinct, weakly sclerotized, opening into ill-defined 1994 Nigeria
Nanniella chalybea Reuter, 1904* tropical Africa Anapomella Putshkov, 1961: 25 (syn. by Namyatova
Nanniella gracilis Linnavuori, 1975 Sudan; & Konstantinov, 2009)
Cameroon Oraniella Reuter, 1894: 138 (syn. by Namyatova &
Nanniella pallidiceps Linnavuori, 1973 Zaire Konstantinov, 2009)
Nanniella palustris Linnavuori, 1975* Equatorial
Africa Diagnosis: Orthocephalus can be recognized by the
Nanniella reuteri Poppius, 1914 south-east Africa following combination of characters: head transverse
with frons flat to moderately convex; eyes sessile; AI
short with two or three mesial spine-like setae,
Biology and host plant associations: No host informa-
dorsum impunctate, usually covered with robust,
tion has been collected for this genus. Nanniella
bristle-like setae, apical portion of ductus seminis
alkithoe has been collected at light in savannah
bowl-shaped, strongly dilated and sclerotized, apically
forest; Nan. gracilis and Nan. chalybea have both
funnel-shaped and dorsoventrally compressed, with
been found in mountain meadows, with the latter also
distinct scale-like sculpturing.
collected in clearings, rain and savannah forests; and
Nan. palustris is known from swampy meadows and
Redescription: Coloration (Fig. 4): body dark brown to
moist clearings in rain and savannah forests, and is
black, sometimes with yellow to reddish-brown
common across western Africa (Linnavuori, 1994).
colouring, particularly on legs; hemelytra of brac-
hypterous morphs sometimes pale tan and brown.
Remarks: Our phylogenetic analysis confirms the Surface and vestiture (Figs 4, 45A–H): dorsum
sister taxon relationship of Nanniella and Acratheus. impunctate, pronotum, scutellum, and hemelytron
As discussed above, both share many similarities, sometimes rugulose; body clothed with long, black,
notably the dense, elongate punctures covering the bristle-like semi-erect setae; AI with two or three
body, the structure of the MTG external efferent spines; femora sometimes with a few spines, tibiae
system, and the male and female genitalia. This rede- strongly spinose; some species also densely clothed in
scription is based on examination of Nan. chalybea white, scale-like setae. Structure: degree of wing poly-
and Nan. palustris, as well as review of the literature morphism varies across species, both sexes mac-
(Linnavuori, 1994). ropterous or brachypterous. Head (Figs 4, 45A–C):
transverse, slightly broader than anterior of prono-
tum, always narrower than posterior of pronotum in
ORTHOCEPHALUS FIEBER (FIGS 4, 45–46) macropterous males; posterior margin of vertex
Orthocephalus Fieber, 1858: 316 (gen. nov.; type rounded to weakly carinate; frons flat to moderately
species: Lygaeus brevis Panzer, 1798 by subsequent convex, steeply declivent; gena height slightly greater
designation Reuter, 1888: 76); Fieber, 1858: 316 (gen. than eye height; eyes round, substylate. Labium
nov., key); Fieber, 1860b: 291, 1861 (key, descr.); (Fig. 45C): variable, reaching from before procoxae to
Douglas & Scott, 1865: 429 (descr.); Thompson, 1871: slightly beyond metacoxae; LI short and swollen.
432 (key); Reuter, 1875b: 24 (key); Reuter, 1875a 1:86; Antennae (Figs 4, 45A–C): insertion close to and in
2:93 (key, descr.); Saunders, 1875: 289 (key); Pro- line with lower margin of eye; thin and cylindrical,
vancher, 1887: 136 (descr.); Reuter, 1891: 43, 158 (key, shorter than body length; AI sometimes somewhat
descr.); Saunders, 1892: 269 (key, descr.); Hueber, swollen, short, sometimes slightly longer than eye
1906: 3, 15 (key, descr.); Kirkaldy, 1906: 131 (cat.); height; AII sometimes apically clavate; AIII and AIV
Oshanin, 1910: 791 (cat.); Reuter, 1910: 148 (cat.); thin. Thorax (Figs 4, 45A–D): pronotum trapezoidal,
Van Duzee, 1914: 389 (cat.); Knight, 1923: 498 (key); in macropterous morphs declivent with thin collar, in
Blatchley, 1926: 804 (descr.); Stichel, 1933: 235 (key); brachypterous morphs sometimes more rectangular
Hedicke, 1935: 59 (key); Knight, 1941: 75, 81 (key); and flat without collar; callosite region present but
Hsiao, 1942: 253 (key); China, 1943: 267 (cat.); Slater, sometimes ill-defined, lateral margins rounded in
1950: 46 (female genitalia); Kiritshenko, 1951: 127 brachypterous morphs, posterior margin straight to
(key); Wagner, 1952: 96, 103 (key, descr.); Carvalho & weakly concave; mesoscutum sometimes concealed by
Leston, 1952: 245 (key, fig.); Carvalho, 1952: 74 (cat.); pronotum in brachypterous morphs; scutellum flat to
Carvalho, 1955: 67 (key); Carvalho, 1958: 22 (cat.); somewhat swollen; metathoracic spiracle conspicuous
Southwood & Leston, 1959: 247 (key); Wagner, 1961: and exposed, often large, narrowly surrounded with
50 (diag., key); Kerzhner, 1964a: 966 (diag., key); evaporative bodies that extend up to dorsolateral
Wagner & Weber, 1964: 267 (descr., key); Wagner, margin of metathorax; MTG efferent system swollen
1973: 41 (descr., key); Schuh, 1995: 63 (world cat.); and triangular, extending as high as base of meso-
Namyatova & Konstantinov, 2009 (generic revision, coxae, ostiole vertical, opens laterally, peritreme oval,
phylogeny). extending posterodorsally from ostiole. Hemelytra
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 46A–D): pygophore conical, posterior margin weakly
divisions present, cuneus long and narrow, membrane asymmetrical, sometimes with thin apical serrate
with two cells; brachypterous – membrane absent, all flange (e.g. Orthocephalus modarresi); parameres of
remaining divisions faint but present, cuneal fracture roughly equal length; left paramere L-shaped, some-
very short, posterior margin rounded. Legs (Figs 4, times with pronounced sensory lobe, apex of apophy-
45E): metafemora moderately swollen; pretarsi sis hooked, sometimes bifid; right paramere with
without fleshy pulvilli. Male genitalia (Figs 45G, H, spoon-like apex deflected laterally, with small apical
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 both portions weakly sclerotized; posterior wall of
ribbing, secondary gonopore moderately sclerotized, bursa copulatrix a simple, undivided, weakly sclero-
basally constricted to form dorsoventral bowl shape, tized plate; vestibulum symmetrical, margins weakly
apex dorsoventrally compressed and laterally sclerotized and unmodified.
expanded, with scale-like sculpturing primarily on
ventral wall, dorsal wall longer than ventral wall;
endosoma usually with two or three spinose spicules, Diversity and distribution: Orthocephalus includes 23
rarely with none. Female genitalia (Fig. 46E, F): scle- species and has a Holarctic distribution. One
rotized rings widely separated, subtriangular, lateral species, Orthocephalus coriaceus, is found both in
and medial margins of DLP adjacent to sclerotized Europe and eastern North America, whereas all
rings weakly sclerotized and upturned; VLP divided other species are restricted to the Palaearctic
into anterior band attached to rami and medial plate, region.
Included species: Orthocephalus arnoldii (Putshkov, (Reuter, 1891) and Salix sp. (Salicaceae) (Namyatova
1961)* Kazakhstan; Ukraine & Konstantinov, 2009) (Table 1).
Orthocephalus bivittatus Fieber, 1864 Turkey;
Turkestan, East Asia (USSR) Remarks: Orthocephalus is superficially similar to
Orthocephalus brevis (Panzer, 1798)* Palaearctic Dasyscytus, Pachytomella, and Anapus, but can be
Orthocephalus championi Saunders, 1894 Corsica readily distinguished by the unique structure of the
Orthocephalus coriaceus (Fabricius, 1777)* secondary gonopore. In a recent revision of Ortho-
Europe; North America cephalus, Anapomella Putshkov and Oraniella Reuter
Orthocephalus fulvipes Reuter, 1904* Mediterra- were designated junior synonyms of Orthocephalus,
nean; Asia Minor based primarily on the secondary gonopore structure
Orthocephalus funestus Jakovlev, 1881 China; (Namyatova & Konstantinov, 2009). This was sup-
Siberia; north-east Asia; Vladivostok ported through a phylogenetic analysis, which also
Orthocephalus medvedevi Kiritshenko, 1951 proposed a sister relationship between Orthocephalus
Ukraine and Pachytomella. Our phylogeny supports this sister
Orthocephalus melas Seidenstücker, 1962 Turkey taxon relationship as well, based on two synapomor-
Orthocephalus minimus Drapolyuk & Kerzhner, phies (69-1: opening of secondary gonopore slit-like;
2000 Kazakhstan 83-1: DLP with pair of sickle-shaped sclerites adja-
Orthocephalus modarresi Linnavuori, 1997 Iran cent to sclerotized rings). Interestingly, although we
Orthocephalus proserpinae (Mulsant & Rey, incorporated the coding of Namyatova & Konstanti-
1852) Mediterranean nov (2009) into our analysis, only the slit-shaped
Orthocephalus putshkovi (Namyatova & Konstanti- secondary gonopore opening was found to be synapo-
nov) Ukraine; Kazakhstan morphic in both analyses. This appears to be the
Orthocephalus rhyparopus Fieber, 1864 south result of differences in taxon sampling, as Namyatova
Russia & Konstantinov (2009) sampled all Orthocephalus,
Orthocephalus saltator (Hahn, 1835) Holarctic not all of which possess paired sickle-shaped sclerites
Orthocephalus scorzonerae Drapolyuk & Kerzhner, on the DLP.
2000 Uzbekistan; Turkmenistan; Kazakhstan
Orthocephalus sefrensis Reuter, 1895 Algeria
Orthocephalus solidus (Seidenstücker, 1971
PACHYTOMELLA REUTER, 1890 (FIGS 4, 47–48)
Turkey Pachytoma Costa, 1842: 288 [gen. nov.; type species:
Orthocephalus styx Reuter, 1908 Turkey; Russia; Pachytoma minor Costa, 1842, =Phytocoris passerinii
Kazakhstan Costa, 1842 by monotypy; junior homonym of Pachy-
Orthocephalus tibialis (Reuter, 1894) Algeria; toma Svains, 1840 (Mollusca)].
Morocco; Tunisia Pachytomella Reuter, 1890: 253 (new name for
Orthocephalus tristis (Reuter, 1894) Algeria Pachytoma Costa, 1842); Reuter, 1891: 37, 158 (key,
Orthocephalus turkmenicus Namyatova & Konstan- descr.); Hueber, 1906: 3, 27 (key, descr.); Oshanin,
tinov, 2009 Turkmenistan; Iran 1910: 796 (cat.); Reuter, 1910: 148 (cat.); Stichel,
Orthocephalus vittipennis (Herrich-Schäeffer, 1933: 235 (key); Hedicke, 1935: 58 (key); China, 1943:
1835) Palaearctic 267 (cat.); Wagner, 1952: 96, 101 (key, descr.); Car-
valho & Leston, 1952: 245 (key, fig.); Carvalho, 1952:
Biology and host plant associations: Host records 74 (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 27
exist for 13 species, all of which have been collected (cat.); Carvalho, 1958: 27 (cat.); Wagner, 1961: 49
mainly on asterids in the family Asteraceae, with a (diag., key); Wagner & Weber, 1964: 265 (syn., descr.,
single species (O. brevis) known only from the Cam- key); Wagner, 1973: 32 (descr., key); Schuh, 1995: 67
panulaceae (Hoberlandt, 1963; Göllner-Scheiding, (world cat.).
1972; Wagner, 1973). In addition to feeding on
asterids, three species have also been collected off Diagnosis: Recognized by the following combination of
rosids, with O. coriaceus found on oak (Fagaceae) characters: head transverse, posterior margin wrap-
(Ehanno, 1960, 1965), O. vittipenis on Ononis spinosa ping around pronotum; hemelytra parallel-sided;
(Fabaceae) (Göllner-Scheiding, 1972), and O. saltator MTG obsolete.
on Spiraea sp. (Namyatova & Konstantinov, 2009).
Orthocephalus saltator has also been collected on Redescription: Sexually dimorphic, with males mac-
unidentified grass (Poaceae) (Ehanno, 1960; Gravest- ropterous or brachypterous and females brachypter-
ein, 1978), several species of Fabaceae (Reuter, 1891; ous. Length = 1.5–4 mm. Coloration (Fig. 4): mostly
Saunders, 1892; Tamanini, 1981; Seidenstücker, dark brown to black, sometimes with the hemelytra
1959), as well as Thymus serpyllum (Lamiaceae) partly to mostly pale, legs usually dark, sometimes
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 ing triangular from above in some species; slightly
vestiture (Figs 4, 47A–H): impunctate, hemelytra and broader than anterior of pronotum, much broader
posterior of pronotum rugulose. Clothed in long, than tall; genae height greater than eye height; pos-
decumbent, simple setae; setae on antennae and legs terior margin of vertex carinate; frons steep; eyes
spinose and semi-erect interspersed with longer large and round, deflected caudally and touching
spines; setae on pygophore long and directed caudally. pronotum; buccula thin; labium (Fig. 47A): LI short
Structure: head (Figs 4, 47A–C): transverse, appear- and broad. Antennae (Figs 4, 47A–C): insertion adja-
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; ribbing, subapically sclerotized and lacking ribbing
AI as long as eye height, somewhat thicker than AII. until secondary gonopore; secondary gonopore
Thorax (Figs 4, 47A, B, D, E): pronotum trapezoidal, roughly circular, usually dorsoventrally compressed
becoming subrectangular in brachypterous morphs, (less so in Pachytomella passerini) with distinct scale-
short, collar absent, callosite region weakly defined in like texturing; endosoma with or without one or
macropterous morphs, obsolete in brachypterous two broad, sclerotized spicules. Female genitalia
females, lateral margins rounded, posterior margin (Fig. 48E, F): sclerotized rings moderately sized, rect-
approximately equal to head width, straight to angular, widely separated, lateral margins and adja-
concave; mesoscutum visible in macropterous males; cent portion of DLP distinctly curved upwards;
scutellum broad in brachypterous morphs; metatho- anterolateral margins of VLP sclerotized; posterior
racic spiracle small and narrow; MTG external effer- wall undivided and plate-like, mostly membranous
ent system obsolete. Hemelytra (Fig. 4): macropterous with weak sclerotization along anterior margin;
males – elongate, parallel-sided; claval commissure opening to vestibulum symmetrical, with lateral
elongate; cuneus narrow and short; membrane with margins weakly sclerotized and unmodified.
two cells, extends beyond abdomen. Brachypterous –
with or without distinct clavus; without cuneus or Diversity and distribution: Pachytomella is a Euro-
membrane; extending partway over abdominal tergite pean genus of six species, which are almost exclu-
VI. Legs (Figs 4, 47F): short; metafemora incrassate sively found in the Mediterranean region.
in females; pretarsi without pulvilli. Abdomen
(Fig. 4): elongate-oval in males, broader and pear- Included species: Pachytomella alutacea (Puton,
shaped in females. Male genitalia (Figs 47G, H, 48A– 1874) Spain
D): pygophore broad and conical; posterior margin Pachytomella cursitans Reuter, 1905b Spain
weakly concave beneath left paramere; parameres Pachytomella doriae (Reuter, 1884)* Spain;
long and slender; left paramere narrow, sensory lobe Tunisia
small or absent, apical apophysis sharply curved or Pachytomella parallela (Meyer-Dür, 1843)*
angled, apex sometimes broadly hooked; right central and north Europe
paramere sometimes projecting from pygophore, Pachytomella passerinii (Costa, 1842)*
apical club elongate, weakly laterally deflected; phal- Mediterranean
lotheca short, basally broad and weakly laterally con- Pachytomella phoenicea (Horváth, 1884) eastern
stricted; ductus seminis attenuate, with flexible Mediterranean
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 Remarks: Within the Halticini, Pachytomella shares
biological records have been published for this genus. with Chorosomella, Plagiotylus, and Strongylocoris
Two species for which host records exist are associ- extreme reduction of the MTG, which based on our
ated with high altitude: Pa. alutacea is found in high phylogenetic analysis, has occurred multiple times
mountains under Thymus spp. and other plants, (character 23-1). As discussed above, Pachytomella is
whereas Pa. parallela is found in mountain meadows sister taxon to Orthocephalus, a relationship sup-
on Potentilla spp., with adults present from June to ported both in our analysis and that of Namyatova &
August (Wagner, 1973). Pachytomella passerini has Konstantinov (2009).
been collected on Potentilla sylvestris (Rosaceae) All three genera are easily distinguished by
(Kerzhner, 1964a) and Thapsia garganica (Apiaceae) their coloration, body shape, and male and female
(Wagner, 1973). This species is associated with arid genitalia.
areas, with adults present from May to August.
Pachytomella passerini overwinter as eggs (Wagner,
1973). Pachytomella doriae has been collected on PIEZOCRANUM HORVÁTH (FIGS 4, 49)
Plantago albicans (Plantaginaceae), with adults Piezocranum Horváth, 1877: 92 (gen. nov.; type
present in May (Wagner, 1973) (Table 1). species: Piezocranum simulans Horváth, 1877 by
monotypy); Atkinson, 1890: 120 (cat.); Reuter, 1891: short, somewhat longer in brachypterous morphs;
33, 157 (descr., key); Hueber, 1906: 3 (key); Kirkaldy, pretarsi without fleshy pulvilli. Abdomen (Fig. 4):
1906: 130 (cat.); Reuter, 1910: 148 (cat.); Oshanin, narrow in macropterous individuals, broader in
1910: 798 (cat.); Stichel, 1933: 235 (key); Kiritshenko, females, in brachypterous females pear-shaped. Male
1951: 127 (key); Wagner, 1952: 96, 101 (key, descr.); genitalia (Fig. 49A–C): pygophore conical; parameres
Carvalho, 1952: 74; Carvalho, 1955: 67 (key); Car- roughly equal in length; left paramere L-shaped,
valho, 1958: 29 (cat.); Kerzhner, 1964a: 967 (diag., sensory lobe broad, apophysis slender and apically
key); Wagner & Weber, 1964: 264 (syn., diag., key); hooked; right paramere slender basally, apical club
Wagner, 1973: 28 (descr., key); Schuh, 1995: 68 (world elongate, weakly deflected laterally, inner margin
cat.). sinuate; phallotheca unmodified; ductus seminis con-
Lamprella Reuter, 1890: 253 (gen. nov.) – (syn. by stricted, with flexible ribbing; secondary gonopore
Horváth, 1889: 327); Lamprella Carvalho, 1958: 29 elongate, incompletely sclerotized, dorsoventrally
(cat.). compressed, apically with scale-like texturing; endo-
soma simple and broad, without spicules. Female
Diagnosis: Recognized by the following combination of genitalia (Fig. 49D–F): sclerotized rings widely sepa-
characters: macropterous individuals elongate, with rated, ovate, weakly sclerotized; lateral margins of
transverse depression spanning vertex and frons; DLP upturned and sclerotized, closely appressed to
endosoma without spicules; DLP divided into weakly rings, remainder of DLP and VLP membranous;
sclerotized lateral and medial plates, with paired posterior wall mostly membranous, with faintly
sclerotized strips adjacent to medial margin of scle- sclerotized band along posterior margin; opening
rotized rings. to vestibulum symmetrical, membranous, and
unmodified.
Redescription: Sexually dimorphic, with males mac-
ropterous and females macropterous or brachypter-
Diversity and distribution: Piezocranum is composed
ous. Coloration (Fig. 4): Male reddish-brown, paler on
of three species, all of which are found almost exclu-
hemelytron; female glossy black. Surface and vesti-
sively in the Mediterranean region, with the range of
ture (Fig. 4): head smooth and shining, pronotum
Piezocranum simulans also extending into the former
rugulose, especially along posterior margin, hem-
Yugoslavia, Bulgaria, and Romania.
elytra of female sometimes glossy with deep punc-
tures. Body of male clothed in golden simple setae;
females less setaceous. Structure: head (Fig. 4): trans- Included species: Piezocranum corvinum Puton,
verse; usually wider than tall; macropterous morphs 1895 Syria; Turkey
with deep transverse depression spanning across Piezocranum seminulum Horváth, 1898 Spain
vertex and frons, brachypterous females with at most Piezocranum simulans Horváth, 1877* southern
a shallow depression; posterior margin of vertex Europe, Macedonia
raised and carinate; frons broadly rounded in brac-
hypterous females; gena height two to three times eye Biology and host plant associations: No biological or
height; labium: LI swollen, short. Antennae (Fig. 4): host plant information has been recorded for this
insertion below or in line with lower margin of eye; genus.
shorter than body, slender; length of AI approximately
equal to eye height. Thorax (Fig. 4): pronotum trap-
ezoidal in macropterous morphs, subrectangular Remarks: This redescription is based on a review of
in brachypterous females, callosite region obsolete the literature, particularly Wagner (1973), and exami-
to weakly defined in both sexes, lateral margins nation of Pi. simulans. Our phylogeny places
rounded, humeral angles rounded, posterior margin Piezocranum as sister to Dasyscytus, although this
concave, more deeply in female; mesoscutum visible relationship is not well supported.
from above in macropterous individuals; metathoracic
spiracle tear-shaped and exposed, without evapora-
tive bodies; MTG external efferent system swollen, PLAGIOTYLUS SCOTT (FIGS 4, 50–51)
angled posterodorsally, ostiole ventrolateral, peri- Plagiotylus Scott, 1874: 272 (gen. nov. type species:
treme tongue-shaped. Hemelytra (Fig. 4): macropter- Plagiotylus maculatus Scott, 1874 by monotypy);
ous – parallel-sided, elongate, extending beyond apex Atkinson, 1890: 125 (cat.); Reuter, 1891: 98, 160
of abdomen; cuneus elongate; membrane with two (descr., key); Hueber, 1906: 5 (key); Kirkaldy, 1906:
cells. Brachypterous females – undivided, posteriorly 131 (cat.); Oshanin, 1910: 783 (cat.); Reuter, 1910: 148
almost straight to subtriangular, sometimes extend- (cat.); Stichel, 1933: 235 (key); Carvalho, 1952: 74
ing partly over abdominal tergite VI. Legs (Fig. 4): (cat.); Carvalho, 1955: 67 (key); Carvalho, 1958: 30
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); pale setae); antennae and legs spinose, spines longest
Wagner, 1973: 100 (descr., key); Schuh, 1995: 69 on AI and tibiae. Structure: male elongate and mac-
(world cat.). ropterous, female oval, coleopteroid. Head (Figs 4,
50A–C): transverse, slightly broader than anterior
Diagnosis: This genus is distinguished from all other margin of pronotum; approximately as tall as broad,
Halticini by the following combination of characters: genae height greater than eye height; posterior
glossy surface; uniform pale green and yellow colora- margin of vertex straight, weakly dorsally rounded;
tion; MTG obsolete; females with coleopteroid vertex with shallow, narrow, transverse depression,
hemelytra. posterior margin weakly upturned and carinate; frons
broadly medially rounded, sulcate at clypeus. Labium
Redescription: Coloration (Fig. 4): mostly pale green, (Fig. 50A, C): LI short and thick; generally reaching
with yellow, brown, or black markings. Head, prono- meso- or metacoxae. Antennae (Figs 4, 50A–C): adja-
tum, and scutellum sometimes with blackish mark- cent to lower margin of eye; always shorter than body,
ings, antennae often with some brown or black shortest in female; AI slightly swollen, almost twice
apically, hemelytra sometimes reddened, legs gener- length of eye height. Thorax (Figs 4, 50A–D): prono-
ally pale. Surface and vestiture (Figs 4, 50A–F): body tum trapezoidal and sloping in males, shorter, sub-
shining and impunctate. Body clothed in long, dark rectangular and flat in females, collar present in
simple setae (Plagiotylus dispar with both dark and males, thin and flat; callosite region weakly defined,
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 broad, apically tapered, with small lateral flanges,
weakly medially concave; mesoscutum visible in both apicodorsally sulcate and weakly sclerotized; left
sexes; metathoracic spiracle elongate-oval, without paramere shorter than right, L-shaped, with broad
evaporative bodies; MTG obsolete. Hemelytra (Figs 4, sensory lobe and narrow apical apophysis, apex typi-
50A): macropterous male – elongate, lateral margins cally weakly hooked; right paramere much longer
subparallel to apex of cuneus; clavus apically expand- than left, apical club weakly laterally deflected,
ing; embolium narrow, R + M vein running nearly to narrow, weakly concave, tapered at apex; ductus
cuneus; cuneus elongate and narrow; membrane with seminis short, thick, with flexible ribbing; secondary
two cells. Coleopteroid female – clavus and R + M gonopore elongate and scoop-shaped with prised oper-
vein faint, cuneus and membrane absent. Legs culum, opens ventrally, apically dorsoventrally com-
(Figs 4, 50E): slightly longer in males; metafemora pressed, with distinct scale-like texture; endosoma
somewhat swollen in females; pretarsi without fleshy simple, without spicules, in P. maculatus (and possi-
pulvilli. Abdomen (Figs 4, 50A): male: elongate, bly others) with weakly sclerotized fields of spines.
parallel-sided to pygophore. Female: broadly Female genitalia (Fig. 51E, F): sclerotized rings ovate,
elongate-oval. Male genitalia (Figs 50A, F, 51A–D): transverse, subcontiguous, weakly sclerotized, most
pygophore conical, genital opening large; phallotheca strongly sclerotized laterally, less so towards midline,
lateral margins and adjacent portion of DLP strongly 1952: 74 (cat.); Carvalho, 1955: 67 (key); Carvalho,
upturned; lateral upturned margins of DLP strongly 1958: 31 (cat.); Wagner & Weber, 1964: 275 (descr.);
sclerotized; lateral-most region of VLP joins with rami Wagner, 1973: 54 (descr.).
to form paired, medially projecting, sclerotized pro-
cesses; posterior wall mostly membranous, with Diagnosis: Recognized by the following combination of
weakly sclerotized, transverse, M-shaped process; characters: head transverse, broader than posterior of
opening to vestibulum symmetrical, lateral margins pronotum; pronotum short, trapezoidal, lateral
weakly swollen and weakly sclerotized. margins subparallel; hemelytra with yellow costal
margins.
Diversity and distribution: Plagiotylus includes six
Redescription: Small, males macropterous, females
species, all restricted to the Mediterranean region.
brachypterous and oblong-oval. Coloration (Fig. 4):
mostly dark brown with lighter yellow and yellow-
Included species: Plagiotylus bolivari (Reuter, brown markings. Hemelytra with yellow costal
1880) Spain margins. Surface and vestiture (Figs 4, 52A–H):
Plagiotylus dispar Reuter, 1899* Syria; Israel head smooth, pronotum weakly rugulose. Body with
Plagiotylus maculatus Scott, 1874* Spain semi-erect simple setae, erect and semi-erect spine-
Plagiotylus ruffoi Tamanini, 1960 Italy like setae, and adpressed scale-like setae. Head with
Plagiotylus sahlbergi Reuter, 1901 Algeria; long spine-like setae and decumbent scale-like setae.
Greece; Libya; Tunisia Antennae with semi-reclining spine-like setae, AI
Plagiotylus zorzii Tamanini, 1955 Italy with several long spines. Pronotum and scutellum
with semi-erect simple setae. Setae on thoracic
Biology and host plant associations: The limited pleura flat and scale-like. Hemelytra and abdomen
biology and host information available for Plagiotylus with caudally directed simple and scale-like setae.
comes primarily from Wagner (1973). Plagiotylus Legs with both simple and spine-like setae, spines at
maculatus is said to prefer dry, sunny areas and is apex of femora and on tibiae long and thick. Struc-
found on Teucrium chamaedrys (Lamiaceae). This ture: head (Figs 4, 52A–D): transverse, wider than
species has a single generation per year, overwinters pronotum; vertex flat, posterior margin rounded,
as eggs, with adults present in June and July. Pla- level with pronotum; frons broadly rounded; genae
giotylus ruffoi has been collected at high altitude height approximately twice eye height; eyes large,
(1200–1890 m) in Sicily, on Astragalus siculus round, bulging, substylate; buccula thin. Labium
nebrodensis (Fabaceae), with adults are present in (Fig. 52D): extends to metacoxae; LI enlarged.
July. Tamanini (1960) also reports this species on an Antennae (Figs 4, 52A–D, F): insertion below lower
unidentified species of Prangos (Apiaceae) (Table 1). 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):
Remarks: This redescription is based on examination pronotum trapezoidal, level, very short, collar faintly
of P. dispar and P. maculatus, as well as the litera- visible in males, callosite region bilaterally tumes-
ture. Wagner (1973) provided descriptions, illustra- cent, lateral margins rounded, posterior of humeral
tions, and a key to all species; however only the angles rounded, posterior margin concave; mesoscu-
parameres and aedeagus of Plagiotylus sahlbergi are tum visible in males; scutellum small; metathoracic
illustrated. spiracle prominent and tear-shaped, with evapora-
The relative placement of Plagiotylus differs tive bodies along anterior margin extending dorsally
between our equal weights and implied weights to hemelytra; MTG external efferent system
analyses, and its true relationship within the Halti- rounded, extends dorsally to metathoracic spiracle,
cini remains uncertain. ostiole directed ventrally, peritreme above ostiole,
rounded, surrounded by evaporative bodies. Hem-
elytra (Figs 4, 52A): macropterous male – parallel-
SCHOENOCORIS COSTA (FIGS 4, 52–53) sided; cuneus elongate, narrow; membrane with two
Schoenocoris Reuter, 1890: 253 (gen. nov.; type cells, extends beyond apex of abdomen. Brachypter-
species: Phytocoris flavomarginatus Costa, 1842 by ous female – undivided; posterior margin concave,
monotypy); Reuter, 1891: 64, 159 (descr., key); shortest at claval commissure; approximately half of
Hueber, 1906: 4, 9 (key, descr.); Kirkaldy, 1906: 131 abdomen exposed. Abdomen: narrow, parallel-sided
(cat.); Oshanin, 1910: 789 (cat.); Reuter, 1910: 148 in males, pear-shaped in females. Legs (Fig. 4):
(cat.); Stichel, 1933: 235 (key); Hedicke, 1935: 59 slightly longer in males; metafemora somewhat
(key); Wagner, 1952: 96, 106 (key, descr.); Carvalho, swollen. Male genitalia (Figs 52F–H, 53A–C): pygo-
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 tapering to blunt apex, opening weakly sclerotized
left paramere; both parameres of similar size, with on dorsal margin; ductus seminis relatively short
long, thin base; left paramere L-shaped, sensory lobe with flexible ribbing; secondary gonopore horseshoe-
broad, apophysis tapering to blunt apex; right shaped, strongly sclerotized basally, apically becom-
paramere extending slightly out of pygophore, api- ing indistinct with faint, scale-like texturing;
cally spoon-shaped, apical margin slightly recurved endosoma with two elongate, irregularly dentate, flat
and weakly, medially cleft; phallotheca elongate-oval, spicules, basally joined with small, weakly sclero-
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): sclero- Diversity and distribution: Represented by a single
tized rings small, elongate-oval, transverse, widely species, Schoenocoris is found only in southern
separated, lateral margins weakly upturned, adja- Europe.
cent portion of DLP sclerotized and upturned; DLP
finely textured with minute spines, anteromedially Included species: Schoenocoris flavomarginatus
cleft; VLP strongly sclerotized, forming two rectan- (Costa, A., 1842)* southern Europe
gular lobes abutting anteromedially; rami with small
weakly sclerotized, medially directed lobes caudal to Biology and host plant associations: Schoenocoris
sclerotized rings; posterior wall of bursa copulatrix lives on Juncaceae, with adults present in July and
simple with weak anterolateral sclerotization, medi- August (Wagner, 1973).
ally with bilaterally symmetrical, posteriorly tapered
and diverging strips; opening to vestibulum sym- Remarks: Schoenocoris is most similar to Dimorpho-
metrical, with small sclerotized strips at anterior coris and Compositocoris in general body structure.
margins. The male genitalia are also similar: in all three
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 Schuh, 1995: 70 (world cat.); Drapolyuk & Kerzhner,
possibly medially joined and situated immediately in 1999: 87 (descr.); Simov, 2005: 213 (descr.).
front of the secondary gonopore. Despite these simi-
larities, our phylogeny does not place Schoenocoris Diagnosis: Recognized by the following combination of
with these genera. Additional data will likely be characters: both sexes short and oval, with staphyli-
needed to assess accurately the phylogenetic place- noid hemelytra; mostly black, often with yellow or
ment of this genus. orange-yellow markings; metafemora incrassate;
ductus seminis with flexible ribbing throughout; sec-
ondary gonopore heavily sclerotized, dorsoventrally
SCIRTETELLUS REUTER (FIGS 4, 54–56) compressed, opening narrow with semiclosed opercu-
Scirtetellus Reuter, 1890: 253 (gen. nov.; type species: lum, apically with scale-like texturing; endosoma
Labops brevipennis Reuter, 1879 by monotypy); with one to three spine- or plate-like spicules; poste-
Reuter, 1891: 67, 159 (descr., key); Hueber, 1906: 4 rior wall of bursa copulatrix sclerotized, often with
(key); Kirkaldy, 1906: 131 (cat.); Oshanin, 1910: 789 prominent inter-ramal tumescences.
(cat.); Reuter, 1910: 148 (cat.); Kiritshenko, 1951: 126
(key); Carvalho, 1952: 74 (cat.); Carvalho, 1955: 66 Redescription: Both sexes staphylinoid. Coloration
(key); Carvalho, 1958: 31 (cat.); Kerzhner, 1964a: 964 (Fig. 4): mostly black, almost always with yellow or
(diag.); Medvedeva, 1975: 73 (descr., disc., key); orange-yellow coloration on head, sometimes with
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 angular, collar indistinct, lateral margins rounded,
usually with yellow margins, abdomen sometimes callosite region indistinct to weakly defined, posterior
with two rows of yellow spots, occasionally merged margin straight to weakly medially cleft; mesoscutum
into two lines, legs brown or black. Surface and ves- not visible; scutellum transverse; metathoracic spi-
titure (Figs 4, 54A–H): glossy and impunctate; vertex racle large and round, surrounded by evaporative
with faint wrinkles radiating from centreline; poste- bodies; MTG external efferent system evaporative
rior of pronotum rugulose. Body with thin simple area large and swollen, ostiole opens ventrolaterally,
setae; antenna with spine-like setae, AI with a few peritreme broadly tongue-shaped, orientated verti-
longer spines on inner surface; legs with semi-erect cally above ostiole. Hemelyra (Figs 4, 54A): staphyli-
spines, most dense on tibiae. Structure: head (Figs 4, noid; short and pad-like, undivided, only covering
54A–D): transverse, broader than pronotum, as tall as base of abdomen. Legs: metafemora incrassate and
broad; posterior of vertex medially level with prono- laterally compressed; metatibiae long; pretarsi
tum; eyes globular, substylate, encroaching on prono- without fleshy pulvilli. Abdomen (Fig. 4): broad,
tum; genae height equal to eye height; frons broadly parallel-sided to elongate oval, broader in females.
rounded, steeply sloping; buccula very thin; labium Male genitalia (Figs 54F–H, 55A–C): pygophore
(Fig. 54C, D): reaches to metacoxae; LI short, moder- conical, genital opening large, posterior margin
ately swollen. Antennae (Figs 4, 54A–D): insertion in weakly concave below left paramere; left paramere
line with or just below lower margin of eye; longer L-shaped, shorter than right, sensory lobe small but
than body; AI swollen, slightly longer than eye height. distinct, apical apophysis thin, curved, apically
Thorax (Figs 4, 54A, C, D): pronotum short flat, rect- hooked or bifid; right paramere longer than left, api-
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 (e.g. Scirtetellus gudali) to strongly upturned (e.g.
apophysis; phallotheca sometimes with dense field of Scirtetellus seminitens), often with medial margin
minute spines on outer surface near apex; ductus also upturned; lateral portion of DLP adjacent to
seminis elongate and narrow with flexible ribbing; sclerotized rings weakly to strongly upturned; VLP
secondary gonopore heavily sclerotized, dorsoven- sclerotized, anteromedially divided or whole; lateral-
trally compressed, opening narrow with semi-closed most region of VLP joins with rami below sclerotized
operculum, apically with scale-like texturing; endo- rings to form paired medially directed sclerotized
soma with one to three spine- or plate-like spicules; in projections, sometimes covered with fields of spines,
some cases one spicule is enclosed in a separate converging with posterior margin of the posterior
pocket of the endosoma. Female genitalia (Figs 55D, wall; posterior wall variable, bilaterally sclerotized in
E, 56A–E): sclerotized rings variable, widely sepa- all species examined, sometimes with small, paired
rated, lateral margins sometimes weakly upturned tumescences or small projections, in others with
elaborate inter-ramal tumescences that may be mented. The presence of inter-ramal lobes on the
dentate and covered in dense fields of spines; opening posterior wall (88-0) of some species is shared with
of vestibulum symmetrical, margins weakly sclero- Anapus and Labops, suggesting a possible close affin-
tized and sinuous. ity, as reflected in our phylogeny. Additionally, the
presence of paired sclerotized lobes formed by fusion
Diversity and distribution: Scirtetellus includes 16 of the VLP and the rami (84-1) is also shared by
species in central and north Asia. several species of Scirtetellus and Labops, indicating
that these structures may be functionally linked.
Included species: Scirtetellus alashanensis Nonnaizab
& Bao, 2006 Mongolia
STRONGYLOCORIS BLANCHARD (FIGS 4, 57–58)
Scirtetellus bianchii Medvedeva, 1975 Asiatic
Russia Strongylocoris Blanchard, 1840: 140 (gen. nov.; type
Scirtetellus brachycerus Kerzhner, 1962 Kazakh- species: Strongylocoris leucocephalus Linnaeus, 1758
stan; Asiatic Russia; Mongolia by monotypy); Costa, 1853: 48 (descr.); Puton, 1886:
Scirtetellus brevipennis (Reuter, 1879)* Turkestan 53 (cat.); Atkinson, 1890: 119 (cat.); Reuter, 1891: 26,
Scirtetellus gudali Kiritshenko, 1951* Georgia, 157 (descr., key); Saunders, 1892: 258 (key, descr.);
Russia, Asiatic Russia Hueber, 1906: 2, 32 (key, descr.); Kirkaldy, 1906: 130
Scirtetellus kerzhneri Medvedeva, 1975 Asiatic (cat.); Reuter, 1910: 148 (cat.); Oshanin, 1910: 799
Russia (cat.); Van Duzee, 1916: 211 (key); Van Duzee, 1917:
Scirtetellus maculiventris Kiritshenko, 1952 276 (cat.); Van Duzee, 1921: 135 (key spp.); Knight,
Asiatic Russia; Tajikistan 1923: 498 (key); Blatchley, 1926: 803 (key spp.);
Scirtetellus micans Medvedeva, 1975 Asiatic Stichel, 1933: 235 (key); Hedicke, 1935: 58 (key);
Russia Knight, 1938: 1 (note); Knight, 1941: 75, 78 (key, key
Scirtetellus mongolicus Drapolyuk & Kerzhner, spp.); Hsiao, 1942: 254 (key); China, 1943: 267 (cat.);
1999 Mongolia Froeschner, 1949: 139 (key spp.); Slater, 1950: 52
Scirtetellus obscurus Medvedeva, 1975 Asiatic (female genitalia); Kiritshenko, 1951: 127 (key);
Russia Wagner, 1952: 95, 99 (key, descr.); Carvalho, 1952: 74
Scirtetellus pallidus Medvedeva, 1975 Asiatic (cat.); Carvalho, 1955: 68 (key); Wagner, 1956: 277
Russia (descr., disc. of generic limits); Carvalho, 1958: 32
Scirtetellus petrovi Simov, 2005* Pakistan (cat.); Southwood & Leston, 1959: 246 (key); Wagner,
Scirtetellus schamili Kiritshenko, 1951 Azer- 1961: 49 (key, diag.); Kerzhner, 1964a: 967 (diag.,
baijan; Asiatic Russia key); Wagner & Weber, 1964: 258 (syn., descr., key);
Scirtetellus seminitens Horváth, 1904* Turkestan; Wagner, 1973: 18 (descr., key); Schuh, 1995: 71 (cat.).
Asiatic Russia Stiphrosomus Fieber, 1858: 312 [gen. (syn. by
Scirtetellus variabilis Medvedeva, 1975 Asiatic Reuter, 1888: 284); Carvalho, 1958: 33 (cat.)].
Russia Stiphrosoma Fieber, 1861: 280 (emendation of
Scirtetellus vittatus Kiritshenko, 1951 Russia; Stiphrosomus Fieber, 1858).
Asiatic Russia
Diagnosis: Characterized by the following combina-
Biology and host plant associations: Scirtetellus is a tion of characters: head broad and short, posterior
polyphagous genus, with most species known from margin closely adpressed to pronotum; pronotum
high mountainous regions of central Asia, in alpine broad, with posterior margin weakly declivent and
and subalpine meadows from 1600 to 4800 m convex; pronotum and hemelytra punctate; bursa
(Medvedeva, 1975). Species are found on various copulatrix complex (Fig. 58G).
plants of the families Asteraceae, Ranunculaceae, and
Poaceae (Medvedeva, 1975). Kirishenko (1952) stated Redescription: Broad and oval, both sexes macropter-
that Scirtetellus are zoogeographically and ecologi- ous, in some species male longer and more slender
cally most similar to Myrmecophyes (Table 1). than female. Coloration (Fig. 4): variable, often black,
brown, or tinged red or blue, often with yellow or
Remarks: The most comprehensive treatment of Scir- yellow-brown markings. Legs sometimes concolour-
tetellus is that of Medevedva (1975, English transla- ous, sometimes partly yellow, yellow-brown, or red.
tion 1977), which includes illustrations of male Surface and vestiture (Figs 4, 57A–G): body clothed in
genitalia and a key to all but the most recently dark or pale simple setae; often with undercoating of
described species, Sc. alashanensis Nonnaizab & Bao short setae interspersed with longer setae; antennae
(2006) and Sc. petrovi Simov (2005). Until now and legs with semi-erect spines. Head glossy, some-
however, the female genitalia have never been docu- times finely rugulose; pronotum and hemelytron
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 bilater- pronotum or nearly so; eye short and tall; head height
ally impunctate; scutellum sometimes rugulose or about two times eye height; frons medially rounded,
punctuate; remainder of body impunctate. Structure: laterally shallowly depressed; base of clypeus not
head (Figs 4, 57A–C): transverse, slightly broader strongly delimited from frons; bucculae very thin.
than anterior of pronotum, broader than tall; vertex Labium (Fig. 57C): LI thick, length approximately
with transverse bisinuate depression, posterior equal to genae height. Antennae (Figs 4, 57A): inser-
margin slightly concave, carinate, generally touching tion well below eyes; short, less than half body length;
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 vein long and close to costal margin; cuneus sort and
(Figs 4, 57A–D): pronotum trapezoidal, broad, collar broad; membrane short, with two cells; membrane
obsolete; callosite region weakly defined, humeral and cuneus often steeply declivent. Legs (Figs 4, 57E):
angles smoothly rounded, posterior margin weakly short; pretarsi without pulvilli. Male genitalia
declivent, rounded or straight; mesoscutum not (Figs 57F, G, 58A–C): pygophore conical, basally
visible; scutellum small, weakly convex; metathoracic broad; both parameres extending somewhat from
spiracle small and tear shaped, surrounded with thin pygophore; left paramere L-shaped, sensory lobe
strip of evaporative bodies; MTG obsolete. Hemelytra broad and rounded, apophysis curved, sometimes
(Fig. 4): broad, lateral margin evenly rounded; R + M twisted, apically hooked; right paramere extremely
long, apically spoon-shaped; phallotheca short, thick, Biology and host plant associations: Host species are
abruptly narrowing from midpoint to apex (in recorded for nine species of Strongylocoris (Table 1).
Strongylocoris leucocephalus apical portion com- Virtually all are associated with asterids, whereas a
pressed from all sides to form dorsal, ventral, and single species, St. luridus, has been collected on the
lateral carinas, dorsally with scale-like texturing); rosid species Calicotome villosa (Fabaceae) (Pericart,
ductus seminis short and thick, with flexible ribbing; 1965).
secondary gonopore elongate and cupped, with prised
operculum opening ventrally; endosoma simple, in at Remarks: Three species of Strongylocoris were dis-
least one species (St. leucocephalus) with three or four sected in this study (St. leucocephalus, St. luridus,
minute sclerotized spicules. Female genitalia and St. steganoides). In all three the female genitalia
(Fig. 57D, E): structure of bursa copulatrix complex; bear the same diagnostic characters, most notably the
sclerotized rings laterally and posteriorly strongly unique structure of the bursa copulatrix. More speci-
sclerotized, becoming indistinct anteriorly, lateral and mens must be examined in order to determine the
medial margins strongly upturned; DLP and VLP degree of variation in this character complex across
closely adpressed in parts, sclerotized, portions adja- the genus.
cent to either side of sclerotized rings folded upwards
with rings, medial portion forming a curved, cylindri-
DISCUSSION
cal enclosure; VLP adjacent to rami weakly sclero-
tized; DLP medially forming a weakly sclerotized The Halticini is retained within the subfamily Ortho-
plate; posterior wall simple and membranous, later- tylinae on the basis of the apically convergent lamel-
ally sometimes with fields of spines; opening of ves- late parempodia. This pretarsal character state also
tibulum symmetrical, medial margins of first occurs in the Phylinae: Pilophorini, but the Halticini
gonapophyses moderately swollen, medial margins of lack the phyline-type male genitalia (s-shaped endo-
vestibular opening weakly sclerotized. soma, phallotheca attached directly to the pygophore).
In contrast, the halticine genitalia are similar to
those found in the nominotypical tribe Orthotylini,
Diversity and distribution: Strongylocoris exhibits a with robust, strap-like spicules and an oval secondary
Palaearctic distribution, with most of the 16 species gonopore.
found in the Mediterranean. 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
Included species: Strongylocoris amabilis (Douglas & the morphology of the posterior wall of the bursa
Scott, 1868) Egypt; Israel copulatrix, including the presence of inter-ramal lobes
Strongylocoris atrocoeruleus (Fieber, 1864) south in three halticine genera, which we believe are
Europe homologous to those found in the Orthotylini and the
Strongylocoris cicadifrons Costa, 1853 Europe; Austromirini Carvalho. An absence of inter-ramal
Israel lobes is the norm for most Halticini, and is also
Strongylocoris coerulescens Lindberg, 1940 Morocco observed in the Nichomachini, and the newly erected
Strongylocoris enki Linnavuori, 1984 Iraq monotypic tribe Coridromiini. On the basis of previ-
Strongylocoris erythroleptus Costa, 1853 Italy ous definitions of the Halticini, these observations
Strongylocoris ferreri Ribes & Pagola-Carte, render the tribe as lacking defining synapomorphies.
2007 Spain Schuh & Lattin (1980), in their paper on the first
Strongylocoris franzi Wagner, 1955 Spain Nearctic Myrmecophyes species, made the observation
Strongylocoris leucocephalus (Linnaeus, 1758)* that this species, M. oregonensis, and another halti-
Palaearctic cine, Anapus americanus, possessed jointly symmetri-
Strongylocoris luridus (Fallén, 1807)* Europe cal and non-overlapping sclerotization of the opening
Strongylocoris niger (Herrich-Schäeffer, 1835) of the vestibulum associated with the first gonapo-
Europe physes; they concluded that this character state was
Strongylocoris oberthuri Reuter, 1905c France worthy of broader comparative investigation in terms
Strongylocoris obscurus (Rambur, 1839) of establishing an alternative synapomorphy for the
Mediterranean tribe. Subsequently, Pluot-Sigwalt & Matocq (2006)
Strongylocoris raimondoi Carapezza, 1991 Italy found similar symmetry for additional halticine
Strongylocoris seabrai Schmidt, 1939 Portugal; species. We too have found such symmetry in the
Spain vestibulum of all halticine females examined;
Strongylocoris steganoides (Sahlberg, 1875)* however such symmetry is also present in the Nicho-
Austria; Russia; Sweden machini, the Coridromiini (= Coridromius), and
several Orthotylini, such as the Hadronema group phylogenetic support for all but a few clades suggests
(Forero, 2008) and Harveycapsus dimorpha (Cassis, that future studies may benefit from the inclusion of
Symonds & Tatarnic, 2010), and can thus not be molecular data, in order to ascertain accurately the
considered as unique to the tribe. relationships of highly autapomorphic genera such as
From the present study a single synapomorphy has Chorosomella, for which morphological character
been identified: the presence of elongate left and right homology is difficult to confirm.
paramere bases. This is not unusual in the speciose
family Miridae, where great morphological diversity
exists, and higher taxonomic level synapomorphies ACKNOWLEDGEMENTS
are rare. Our definition of the Halticini retains the We wish to thank the museums mentioned herein for
tribe as a highly diverse suprageneric group with a supplying the material for this work. We are also
number of highly autapomorphic genera (e.g. grateful to the following individuals for providing
Chorosomella, Dicyphopsis). fresh material for study: freshly collected specimens
This study has produced robust definitions of the of Myrmecophyes alboornatus were provided by Fedor
genera, inclusive of information for the male and Konstantinov of the Zoological Institute of Saint
female genitalia, with exemplars illustrated for Petersburg; specimens of Halticus apterus were pro-
nearly all genera. On the basis of external characters vided by Michael D. Schwartz of the Agriculture and
such as head structure, wing polymorphism, the Agri-Food Canada (CNC, Ottawa); Randall T. Schuh
external efferent system of the MTG, and the genita- from the American Museum of Natural History pro-
lia, we have mostly supported the previous generic vided specimens of Compositocoris senecionus;
status of the halticines as summarized in the world Thomas Henry from the U.S. Natural History
catalogue of Schuh (1995). Apart from the removal of Museum and Dimitri Forero (AMNH) provided speci-
Coridromius from the Halticini, the most significant mens of recent collections of Cafayatina altigena from
changes we present are in the nominate genus Hal- Argentina. Additionally, Nikolay Simov provided
ticus, which as previously defined was not monophyl- specimens of Dimorphocoris fuscus from Bulgaria,
etic. Our subdivision of the genus leads to the and the recently described Scirtetellus petrovi from
following outcomes: (1) Halticus has been redefined to Pakistan. We also thank Sue Lindsay (Australian
include only those species, which as a group have a Museum) for providing assistance with the scanning
Holarctic distribution, and are defined by large, electron micrographs of most taxa. This manuscript
transverse, subcontiguous, female sclerotized rings, was significantly improved through comments from
the scoop-shaped secondary gonopore, and vestigial two anonymous reviewers. Lastly, we thank Michael
endosoma; (2) Ectmetopterus is redefined to include Schwartz, who provided the senior author with tre-
east Asian species previously placed within Halticus mendous assistance throughout this project. This
and now recognized by the tripartite left paramere; research was conducted during the course of the
and (3) Microtechnites has been removed from syn- senior author’s PhD, which was funded through the
onymy, with its generic status restored for a group of NSF Planetary Biodiversity Inventories grant DEB-
mostly Neotropical species defined by the multiple 0316495 to R. T. Schuh and G. Cassis.
spicules and the small and widely separated female
sclerotized rings. Based on our phylogenetic analysis
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(Hemipt. Heteropt.). Revue Française d’Entomologie 21:
APPENDIX 1
219–223.
Wagner E. 1955. Bemerkungen zum System der Miridae LIST OF CHARACTERS AND CHARACTER STATES
(Hem. Het.). Deutsche Entomologische Zeitschrift, N. F. 2: (Figures cited refer to examples of structures
230–242. described.)
Wagner E. 1956. La validité de l’espèce Strongylocoris ober-
thuri Reuter. Vie et Milieu 7: 101–103. 1. Coloration: (0) variably coloured, not mostly
Wagner E. 1957. Zwei neue Dimorphocoris-Arten aus black; (1) mostly black or dark brown.
Frankreich (Hem. Het. Miridae). Deutsche Entomologische 2. Texture – head and thorax: (0) without patches of
Zeitschrift, N.F. 4: 223–227. scale-like or honeycombed texturing; (1) with
Wagner E. 1960. Was ist Dasyscytus sordidus Fieber 1864? symmetrical patches of scale-like honeycombed
(Hem. Het. Miridae). Mitteilungen der Deutsche Entomolo- texturing laterally on thorax and dorsally on
gischen Gesellschaft 19: 98–101. head (Figs 15A, C, D, 22E).
3. Punctures – body and hemelytra: (0) impunctate, 19. AI length: (0) less than height of eye (Figs 17B, C,
or with very few punctures; (1) nearly entirely 28B); (1) equal or slightly greater than height of
punctate (Figs 5A–C, E, 43A–C, E). eye (Fig. 37B, C); (2) much longer than height of
4. Rugosity – pronotum: (0) not rugose (Figs 5A, eye (Figs 13A, 33B, C).
20A, 22A, 50B); (1) wholly or partly rugose 20. AIV length: (0) shorter or equal to AIII; (1) longer
(Figs 7A, 11A, 26A, B, 28A, 39A, 52B). than AIII.
5. White scale-like or feather-like setae: (0) absent; 21. Total antennal length: (0) subequal to body
(1) present on head, thorax, hemelytra or legs length; (1) longer than body.
(Fig. 3: Anapomella arnoldi, Dampierella 22. Metathoracic spiracle shape: (0) narrow and slit-
schwartzi; Fig. 17A–D). like; (1) teardrop-shaped (Fig. 39D).
6. Erect or semi-erect bristle-like setae on dorsum: 23. Metathoracic scent gland: (0) with MTG opening,
(0) absent, or with only a few erect or semi-erect peritreme and evaporative area all well developed
bristle-like setae; (1) dorsum clothed in erect or (Figs 11D, E, 39D, E); (1) some or all components
semi-erect dark, bristle-like setae (Fig. 41A: col- of MTG efferent system reduced or obsolete.
oration not visible in scanning electron micro- 24. Metathoracic scent gland – size of evaporative
graph); (2) dorsum clothed in erect or semi-erect area: (0) evaporative area minute or absent
pale, bristle-like setae. (Figs 47C, D, 50C, D, 57C, D); (1) small, occupy-
7. Texture – frons: (0) without radiating folds; (1) ing no greater than half metepimeron (Fig. 5E,
with radiating folds diverging anteriorly F); (2) large, extending nearly to dorsum
(Figs 26B–C, 39B). (Figs 11D, E, 26F).
8. Vertex sinuation (viewed anteriorly): (0) dorsal 25. Metathoracic scent gland – orientation of peri-
margin of vertex appearing flat or convex, but treme: (0) vertical (Fig. 26F); (1) angled somewhat
never concave (Figs 5C, 11B, 20B, 24B, 26C, caudally (Fig. 33C, D); (3) angled back and
37B, 43B, 45B, 57B); (1) concave (Figs 33B, 35A, resting along caudal margin of tergite (Figs 5E, F,
B). 43E, F).
9. Vertex – posterior margin: (0) rounded (Figs 33A, 26. Metathoracic scent gland – peritreme shape: (0)
39A, 54A); (1) sharply edged (Fig. 45A, C);. long and narrow or teardrop-shaped (Figs 5E, F,
10. Head shape as viewed from above: (0) transverse 33D, 39D, 43E, F); (1) broadly elongate-oval or
(much wider than long) (Figs 5A, 57B); (1) elon- round (Figs 37D, E, 52E).
gate (subequal, equal, or longer than wide) 27. Angle of proepisternum (viewed laterally): (0)
(Fig. 15C). surface of proepisternum immediately adjacent to
11. Paired, textured spots on vertex, superficially anterior thoracic margin laterally orientated
similar to ocelli: (0) absent; (1) present (Fig. 52B). (Fig. 33C); (1) surface of proepisternum anteriorly
12. Conical projection of frons: (0) not conically pro- orientated and appressed to posterior of head
jecting; (1) conically projecting (Fig. 13A, B). (Figs 30D, 37C).
13. Head height below eyes: (0) significantly less than 28. Pronotal shape (macropterous): (0) trapezoidal
height of eye; (1) less than or equal to height of (Figs 5A, 17A, 24A); (1) campanulate (Figs 22A,
eye (Figs 13B, 43D); (2) greater than height of eye 33A).
but less than twice height of eye (Figs 37C, 47A); 29. Pronotal collar: (0) absent or vestigial (Fig. 7A);
(3) greater than twice as high as the eye (1) present in at least one sex (Fig. 20A).
(Figs 33B, C, 39C). 30. Posterior margin of pronotum: (0) straight
14. Labial length: (0) reaching procoxa; (1) reaching (Fig. 39A); (1) concave (Fig. 20A); (2) convex
mesocoxa; (2) reaching metacoxa; (3) reaching (Figs 37A, 57A).
abdomen (surpassing metacoxa). 31. Mesoscutum (macropterous): (0) not visible from
15. Angle of clypeus (viewed laterally): (0) appearing above; (1) visible from above.
vertical or nearly so (Figs 24D, 33C); (1) distinctly 32. Mesoscutum (brachypterous): (0) not visible from
recurved and angled posteriorly (Fig. 39C). above; (1) visible from above.
16. Stalked eyes: (0) eyes sessile; (1) eyes stalked 33. Hemelytral membrane cells: (0) two cells; (1)
(Fig. 33A–C). single cell (Figs 17A, 28A); (2) no cells.
17. Antennal fossa: (0) in line with medial portion of 34. Hemelytral polymorphism – male: (0) always or
eye (Fig. 5B); (1) in line with lower margin of eye nearly always macropterous, with all components
(Figs 17B-C, 28B, C); (2) below eye (Fig. 47A). present; (1) polymorphic, with both macropterous
18. AI spines: (0) smooth or with minute spines and reduced-wing morphs; (2) always with
only (Fig. 37A–C); (1) with a few long reduced wings.
spines (Fig. 47A); (2) with multiple long spines 35. Hemelytra - medial fracture (reduced wing): (0)
(Figs 7A–C, E, 26A-E). absent; (1) short (subequal to length of claval
suture); (2) elongate (greater than length of claval 53. Left paramere - sensory lobe: (0) without discern-
suture). ible sensory lobe (Figs 31D, 40B, 48C); (1) sensory
36. Hemelytra- cuneus (reduced-wing): (0) absent; (1) lobe present but poorly developed (Figs 23B, 46C);
present. (2) sensory lobe large and conspicuous (Figs 9B,
37. Hemelytra – degree of female brachyptery: (0) 12B, 27A).
macropterous – all components present; (1) all 54. Left paramere - apophysis: (0) without recurved
components present but membrane reduced to a apical hook or knob (Fig. 6B); (1) with small down-
thin strip; (2) reduced wing (brachypterous, turned hook or knob (Figs 9B, 12B); (2) with
coleopterous, staphylinoidous) (Fig. 7A); (3) bifurcate, hooked tip (Figs 14B, 19B).
micropterous (wing pads) (Fig. 39A). 55. Coupling of aedeagus and left paramere: (0) not
38. Hemelytral length – (reduced-wing female): (0) coupled to form a piercing intromittent organ; (1)
exposing fewer than two tergites; (1) exposing two coupled to form a piercing intromittent organ.
to three tergites; (2) exposing more than three 56. Phallotheca – apex: (0) without laterally projecting
tergites. apical flange; (1) with prominent laterally project-
39. Hemelytra – membrane setae: (0) naked; (1) ing apical flange (Fig. 36A).
covered with dense setae. 57. Phallotheca – apical constriction: (0) not laterally
40. Hemelytra – lateral corial margin: (0) straight or constricted; (1) abruptly laterally constricted
convex; (1) sinuate. beyond base (Fig. 44A).
41. Hind femur thickness – female: (0) thin; (1) incras- 58. Ductus seminis – length and width: (0) relatively
sate. thick and stout, sometimes short (Figs 18C, 31B,
42. Pretarsus – fleshy pulvilli: (0) underside of claws 44A, B, 46A); (1) slender, long, and sinuous, fre-
without fleshy pulvilli (Figs 39F, 45E, 47F); (1) quently convoluted (Figs 9A, 14A, 16A, B, 21A,
underside of claws with fleshy pulvilli (Figs 7F, 49A).
11F, 17H, 24F). 59. Ductus seminis – ribbing: (0) present (Fig. 18C);
43. Foretibiae – shape: (0) not apically swollen; (1) (1) absent (Fig. 44B).
apically swollen and laterally compressed. 60. Secondary gonopore – apical extension: (0) does not
44. Metatibia – spines: (0) without spines; (1) spinose. extend as a sclerotized tube running to the apex
45. Metafemora – spines: (0) absent; (1) present. of the phallotheca; (1) extends as a sclerotized
46. Metafemora – apex: (0) medially and laterally tube to the apex of the phallotheca (Figs 6A, 44A,
without rounded tumescences at apex; (1) medially B).
and laterally with round, smooth, and slightly 61. Secondary gonopore – apical sclerotization: (0)
swollen tumescence at apex. with a short or long sclerotized section beginning
47. Pygophore – posteroventral margin: (0) symmetri- subapical to opening (Figs 8A, 55A); (1) mostly
cal or only weakly asymmetrical (Figs 37G, H, unsclerotized, if sclerotized then only along rim of
39H, 45G, 47H); (1) asymmetrical, with deep secondary gonopore (Figs 16B, 21A).
concavity below left paramere (Figs 7E, 33F–H); 62. Ductus seminis – apical sclerotization: (0) apical
(2) asymmetrical and complex [significantly sclerotized region of the ductus seminis continu-
rotated, armed with spicules, etc. (outgroup ous, unbroken by clear transverse band (Fig. 40A);
only)]. (1) apical sclerotization of the ductus seminis
48. Left and right paramere – base: (0) neither broken by clear transverse band prior to the
paramere with elongate base; (1) both parameres secondary gonopore (Fig. 48B).
with elongate and narrow base (Figs 9B, C, 12B, 63. Ductus seminis – length of apical sclerotization: (0)
C); (2) base or right paramere elongate, that of left apical sclerotized region short, barely extending
paramere much shorter (Figs 18A, B, 29A, B). below the secondary gonopore (Figs 46A, 53A,
49. Right paramere – apex: (0) not clubbed and cupped; 55A); (1) apical sclerotized region of intermediate
(1) clubbed and cupped. length (Figs 18C, 34A, 40A); (2) apical sclerotized
50. Right paramere - length: (0) much shorter than region elongate (Figs 8A, 9A, 27C).
left; (1) slightly shorter to slightly longer than left; 64. Secondary gonopore – width (when viewed from
(2) much longer than left. above): (0) narrow; (1) broad (Fig. 44A).
51. Extension of right paramere: (0) not extending or 65. Secondary gonopore – dorsoventral compression:
only slightly extending from pygophore; (1) (0) not dorsoventrally compressed, more or less
extending considerably from pygophore (Figs 30H, cylindrical; (1) dorsoventrally compressed
33G, H, 47G, H). (Fig. 46B).
52. Left paramere - shape: (0) more or less L-shaped; 66. Secondary gonopore – shape: (0) forming a narrow
(1) trifurcate (Fig. 25B); (2) twisted and elongate scoop (Figs 18C, 31B, C); (1) not forming a narrow
(Fig. 31D). scoop.
67. Secondary gonopore – ventral wall: (0) ventral wall 81. Ventral labiate plate – region bordering rami: (0)
obsolete, not distinctly recurved; (1) ventral wall not forming a sclerotized rim; (1) forming a mod-
distinctly recurved, forming a prised operculum erately to strongly sclerotized rim (Fig. 9E).
(Figs 9A, 51A, 55A, 58C). 82. Inter-ramal bridge: (0) absent; (1) present
68. Secondary gonopore – subapical shape: (0) without (Figs 10A, 34G, 36E, F).
distinct subapical bowl-shaped constriction; (1) 83. Dorsal labiate plate – medial sclerites: (0) absent;
with distinct sub-apical bowl-shaped constriction (1) with paired, sickle-shaped sclerites adjacent to
and funnel-shaped opening (Fig. 46A, B). sclerotized rings.
69. Secondary gonopore – shape of opening: (0) 84. Rami – medial projections: (0) absent; (1) present
opening of secondary gonopore oval; (1) opening of (Figs 34G, 35A).
secondary gonopore slit-like. 85. Posterior wall – sclerotization: (0) wholly or partly
70. Secondary gonopore – texture: (0) smooth; (1) with sclerotized; (1) entirely membranous (Figs 6F,
scale-like texture (Fig. 46B). 14E, 44H).
71. Endosoma: (0) reduced or absent (Fig. 31B, C); (1) 86. Posterior wall – texture: (0) with fields of spines
forming a well-developed membranous sac (Figs 34H, 44H); (1) without fields of spines
(Figs 34A, 36B). (Fig. 6F).
72. Endosoma – spicules: (0) absent; (1) present. 87. Posterior wall – structure: (0) without bilateral
73. Endosoma – spicule location: (0) attached basal pillow-like swellings; (1) with bilateral pillow-like
to secondary gonopore; (1) distal to secondary swellings (Figs 6F, 25E, 44H).
gonopore. 88. Posterior wall – inter-ramal lobes: (0) absent; (1)
74. Endosoma – spicule form: (0) without paired scler- with prominent, paired, projecting spinose or
ites attached to anterior of secondary gonopore; (1) serrate lobes (Figs 8E, 9F, 10B, 34H, 35B, 36G,
with paired paddle-like sclerites extending from 55D, 55C).
anterior of secondary gonopore (Figs 16A, B, 23A, 89. Posterior wall – base of inter-ramal lobes: (0)
53A). paired, hollow inter-ramal lobes narrow and
75. Endosoma – spicule form: (0) when present, simple strongly constricted basally (outgroup only); (1)
(Figs 9A, 12A); (1) spicules, when present complex inter-ramal lobes not narrow and basally
and branching (Figs 29C, D, 38A). constricted.
76. Endosoma – spicule form: (0) without tight bunch 90. Posterior wall – division: (0) distinctly tripartite,
of spinose spicules; (1) with tight bunch of spinose with discrete lateral plates above a smaller medial
spicules (Figs 34A, 36B). plate (outgroup only); (1) occasionally bilaterally
77. Endosoma – fields of spines: (0) without fields of modified, but without discrete lateral and basal
spines; (1) apically with sclerotized fields of spines plates.
(Figs 19A, 51A); (2) apically with one or two dis- 91. Opening to vestibulum- symmetry of first gonapo-
crete, elongate sclerites covered with fields of physes: (0) asymmetrical; (1) symmetrical
spines (Figs 34A, 40A). (Figs 6E, 10C, 25G).
78. Female - sclerotized rings: (0) present; (1) absent. 92. Opening to vestibulum – degree of sclerotization:
79. Female - sclerotized rings: (0) widely separated; (0) entirely membranous (Fig. 14D); (1) with at
(1) medially contiguous or subcontiguous. least some sclerotization (Figs 6E, 10C).
80. Female - sclerotized rings: (0) no larger than
diameter of posterior wall; (1) larger than diam-
eter of posterior wall.
APPENDIX 2
Halticini morphological data matrix.
1 1 1 1 1 1 1 1 1 1 2
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Orthotylus sp. 0 0 0 0 0 0 0 0 0 1 0 0 1 2 0 0 2 0 1 0
Ceratocapsus sp. 0 0 0 0 0 2 0 0 1 0 0 0 0 2 1 0 2 1 0 0
Nichomachus minutus 0 0 0 1 0 0 0 1 1 0 0 0 0 2 1 0 1 0 0 0
Aopolonema princeps 1 0 0 0 ? 0 0 0 1 0 1 0 0 ? 0 0 1 0 ? 0
Lattinova jacki 0 0 0 0 0 0 0 0 0 1 0 0 0 2 0 0 1 1 2 0
Acratheus ocellaris 1 0 1 0 0 0 0 0 1 0 0 0 2 2 0 0 1 0 1 0
Anapus americanus 1 0 0 1 1 0 1 0 0 0 0 0 3 2 1 0 3 2 2 0
Anapus kirshbaumi 1 0 0 1 1 0 0 0 0 0 1 0 3 1 1 0 3 2 2 0
Anapus rugicollis 1 0 0 1 1 0 1 0 0 0 1 0 3 2 1 0 3 2 2 0
Anapus dorsalis 1 0 0 1 0 0 1 0 0 0 1 0 3 2 1 0 3 2 2 0
Barbarosia punctulata 1 0 0 1 0 0 0 0 0 0 0 0 3 1 1 0 3 1 1 0
Chorosomella jakowleffi 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 1 0 2 0
Coridromius variegatus 0 0 0 0 0 0 0 0 1 0 0 0 1 2 0 0 1 0 0 0
Dasyscytus sordidus 1 0 0 1 1 2 0 0 1 0 0 0 2 1 1 0 3 1 1 ?
Dicyphopsis sp. 0 0 0 0 0 0 0 0 1 0 0 0 1 2 1 0 1 1 1 0
Compositocoris senecionus 0 1 0 0 1 1 0 0 0 1 0 0 1 1 0 0 2 2 2 0
Dampierella schwartzi 1 0 0 0 1 0 0 0 1 0 0 0 1 2 1 0 2 1 0 0
Dimorphocoris gracilis 0 1 0 0 1 1 0 1 0 1 0 0 2 2 0 0 3 2 2 0
Dimorphocoris fuscus 0 1 0 0 1 1 0 1 0 1 0 0 3 2 0 0 3 2 2 0
Ectmetopterus micantulus 1 0 0 0 1 0 0 0 1 0 0 0 2 2 0 0 1 1 1 1
Euryopicoris nitidus 1 0 0 1 0 0 1 0 0 0 1 0 3 1 1 0 3 2 2 1
Goodeniaphila cassisi 1 0 0 1 0 0 0 0 1 0 0 0 1 1 1 0 2 0 0 0
Halticus apterus 1 0 0 1 1 0 1 0 1 0 0 0 2 1 0 0 1 1 0 1
Halticus intermedius 1 0 0 1 1 0 1 0 1 0 0 0 2 1 0 0 1 1 1 1
Halticus luteicollis 1 0 0 1 1 0 0 0 1 0 0 0 2 0 0 0 1 1 1 1
Labops burnmeisteri 1 0 0 1 1 2 1 1 0 0 1 0 3 2 0 1 3 2 2 0
Labops utahensis 1 0 0 1 1 2 1 1 0 0 1 0 3 2 0 1 3 2 2 0
Labops sahlbergii 1 0 0 1 1 2 1 1 0 0 1 0 3 2 0 1 3 2 2 0
Labops tumidiferons 1 0 0 1 1 2 1 1 0 0 1 0 3 1 0 1 3 2 2 0
Microtechnites altigena 1 0 0 1 1 0 0 0 1 0 0 0 2 2 0 0 1 0 0 0
Microtechnites bractatus 1 0 0 1 1 0 0 0 1 0 0 0 2 2 0 0 1 0 1 0
Myrmecophyes alboornatus 1 0 0 1 1 0 1 1 0 0 1 0 3 1 1 0 3 1 2 0
Myrmecophyes oregonensis 1 0 0 1 1 0 1 0 0 0 1 0 3 2 1 0 3 1 2 0
Namaquacapsus melanostethoides 1 0 0 0 0 1 0 0 1 0 0 0 2 2 1 0 2 0 0 0
Nanniella palustris 1 0 1 0 0 0 0 0 1 0 1 0 1 1 0 0 1 0 1 0
Orthocephalus arnoldii 1 0 0 1 1 1 0 0 1 0 0 0 2 2 1 0 3 0 0 ?
Orthocephalus brevis 1 0 0 1 1 1 0 0 1 0 1 0 2 1 0 0 3 1 1 0
Pachytomella passerini 1 0 0 1 0 1 0 0 1 0 1 0 2 1 1 0 3 1 1 0
Piezocranum simulans 1 0 0 1 1 1 0 0 1 0 1 0 2 2 1 0 3 0 0 0
Plagiotylus maculatus 0 0 0 0 1 1 0 0 1 0 1 0 2 2 1 0 3 1 2 0
Schoenocoris flavomarginatus 0 0 0 1 1 1 0 0 0 0 1 0 2 2 1 0 3 2 2 0
Scirtetellus seminitens 1 0 0 1 1 1 1 1 0 0 1 0 3 2 1 0 3 2 2 0
Scirtetellus brevipennis 1 0 0 1 1 1 1 1 0 0 1 0 3 2 1 0 3 2 2 0
Strongylocoris leucocephalus 1 0 0 0 0 0 0 0 1 0 1 0 2 1 1 0 3 1 0 0
2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 4 4
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Orthotylus sp. 0 1 0 1 1 1 1 0 0 0 0 - 0 0 - - 0 - 0 0
Ceratocapsus sp. 0 1 0 1 2 0 1 0 1 0 1 - 0 0 - - 0 - 0 0
Nichomachus minutus 0 1 0 1 1 0 1 0 1 1 1 ? 0 0 ? ? ? ? 0 1
Aopolonema princeps 0 1 0 1 1 0 1 0 1 0 1 - 0 0 - - 0 - 0 0
Lattinova jacki 1 1 0 1 1 0 1 1 0 1 1 - 1 0 - - 0 - 0 0
Acratheus ocellaris 1 1 0 1 3 0 0 0 1 1 0 - 0 0 - - 0 - 0 0
Anapus americanus 1 1 0 1 1 1 0 0 0 1 1 0 - 2 0 0 3 2 0 0
Anapus dorsalis 0 1 0 1 1 1 0 - 0 1 - 0 - 1 0 0 2 2 0 0
Barbarosia punctulata 0 1 0 2 1 1 1 - 0 1 - 0 - 1 0 0 2 1 - 0
Chorosomella jakowleffi 1 1 1 0 - - 0 0 0 1 1 1 1 0 1 0 3 2 0 0
Coridromius variegatus 0 0 1 1 3 0 1 0 1 1 1 - 0 0 - - 0 - 0 0
Dasyscytus sordidus ? 1 0 1 1 0 1 0 ? 1 0 0 0 0 0 0 2 2 1 0
Dicyphopsis sp. 1 1 0 1 3 0 0 1 1 1 0 - 0 0 - - 0 - 0 0
APPENDIX 2 Continued
2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 4 4
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Dampierella schwartzi 0 1 0 1 1 1 1 0 0 1 0 - 1 0 - - 0 - 0 0
Ectmetopterus micantulus 1 1 0 1 3 0 0 0 1 0 1 - 0 0 - - 0 - 0 0
Goodeniaphila cassisi 0 1 0 1 1 1 1 0 0 1 0 - 1 0 - - 0 - 0 0
Halticus intermedius 1 1 0 1 1 0 1 0 0 0 0 - 0 0 - - 0 - 0 0
Halticus luteicollis 1 1 0 1 1 1 1 0 1 0 0 - 0 0 1 1 0 - 0 0
Microtechnites altigena 0 1 0 1 1 1 1 0 1 2 0 - 0 0 - - 0 - 0 0
Namaquacapsus melanostethoides 0 1 ? ? ? ? 0 0 1 0 0 ? 0 0 ? ? ? ? 0 0
Nanniella palustris 0 1 0 1 3 0 0 0 1 1 0 - 0 0 - - 0 - 0 0
Orthocephalus arnoldii 0 1 0 1 1 1 0 0 0 0 1 0 0 0 0 0 2 1 0 0
Pachytomella passerini 0 1 1 0 - - 1 0 0 1 1 0 0 0 0 0 2 1 0 0
Plagiotylus maculatus 0 1 1 0 - - 1 0 1 1 1 1 0 0 1 0 2 1 0 0
Scirtetellus seminitens 1 1 0 1 1 0 0 - 0 0 - 0 - 2 0 0 3 2 - 0
Scirtetellus brevipennis 1 1 0 1 1 0 0 - 0 0 - 0 - 2 0 0 3 2 - 0
Strongylocoris leucocephalus 0 1 1 0 - - 1 0 0 2 0 - 0 0 - - 0 - 0 0
4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Orthotylus sp. 0 0 0 1 0 0 2 0 0 1 0 0 1 0 0 0 0 0 0 0
Ceratocapsus sp. 0 1 0 1 0 0 1 0 0 1 1 1 2 2 0 0 0 0 0 0
Nichomachus minutus ? 0 0 1 0 0 1 0 0 0 0 0 2 1 0 0 0 1 1 0
Aopolonema princeps 0 1 0 1 ? 0 2 1 1 1 0 0 0 1 0 0 0 1 0 0
Lattinova jacki 0 0 0 1 1 0 2 0 1 1 0 0 2 0 0 0 0 0 0 0
Acratheus ocellaris 0 0 0 0 0 0 0 1 1 1 1 0 1 0 0 0 1 0 1 1
Anapus dorsalis 1 1 1 1 1 1 1 1 1 2 0 0 2 1 0 0 0 1 0 0
Chorosomella jakowleffi 0 1 0 1 0 0 0 1 1 1 0 0 1 2 0 0 0 1 0 0
Coridromius variegatus 1 1 0 1 0 1 1 0 1 0 0 0 0 0 1 0 0 1 1 0
Dasyscytus sordidus 0 0 0 1 0 ? 0 1 1 1 0 0 2 2 0 0 0 1 0 0
Dicyphopsis sp. 1 0 0 1 0 0 0 1 1 2 0 0 1 1 0 0 0 1 0 0
Halticus intermedius 1 1 0 1 0 1 1 1 1 2 1 2 0 1 0 0 0 0 0 0
Halticus luteicollis 1 1 0 1 0 1 1 1 1 2 1 2 0 1 0 0 0 0 0 0
Namaquacapsus melanostethoides 0 1 0 0 0 0 ? 1 1 2 ? 0 2 2 0 0 0 1 0 0
4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Nanniella palustris 0 0 0 0 0 0 0 1 1 1 0 0 2 2 0 0 1 0 1 1
Orthocephalus arnoldii 1 0 0 1 1 0 ? 1 1 1 0 0 2 1 0 0 0 ? 0 0
Plagiotylus maculatus 1 0 0 1 1 0 0 1 1 2 0 0 2 1 0 0 0 0 0 0
6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 7 8
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Orthotylus sp. 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0
Ceratocapsus sp. 0 0 1 0 0 0 1 0 0 0 0 1 - 0 0 0 0 0 1 0
Nichomachus minutus 0 0 1 0 0 0 0 0 0 1 0 1 - 0 0 0 1 0 1 0
Aopolonema princeps 0 0 2 0 0 0 0 0 0 ? 0 0 0 0 1 0 0 0 0 0
Lattinova jacki 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 1 0 0 0 0 0
Acratheus ocellaris 0 0 - 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0
Anapus dorsalis 0 1 2 0 0 1 1 0 0 0 1 0 1 0 0 0 2 0 0 0
Chorosomella jakowleffi 1 - - 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 1 0
Coridromius variegatus 1 - - 0 0 1 0 0 0 0 0 1 - 0 0 0 0 1 - -
Dasyscytus sordidus 0 0 0 0 1 1 0 0 0 0 1 0 1 0 0 0 1 0 0 0
Dicyphopsis sp. 1 - - 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 1 0
Compositocoris senecionus 1 - - 0 0 1 0 0 0 0 1 0 1 1 0 0 0 0 0 0
Dimorphocoris gracilis 1 - - 0 1 1 0 0 0 1 1 0 1 1 0 0 0 0 0 0
Dimorphocoris fuscus 1 - - 0 1 1 0 0 0 0 1 0 1 1 0 0 0 0 0 0
Halticus apterus 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 0 0 1 1
Halticus intermedius 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 0 0 1 1
Halticus luteicollis 0 0 0 0 0 0 0 0 0 0 0 1 - 0 0 0 0 0 1 1
Namaquacapsus melanostethoides 0 1 1 0 0 1 0 0 0 1 1 1 - 0 0 0 1 ? ? ?
Nanniella palustris 0 0 - 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0
Orthocephalus arnoldii 0 0 0 0 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0
Piezocranum simulans 0 0 0 0 1 1 0 0 0 1 1 1 - 0 0 0 0 0 0 0
Plagiotylus maculatus 0 1 0 0 1 1 1 0 0 1 1 1 - 0 0 0 1 0 1 0
8 8 8 8 8 8 8 8 8 9 9 9
1 2 3 4 5 6 7 8 9 0 1 2
Orthotylus sp. ? 0 0 0 0 0 0 0 0 0 0 1
Ceratocapsus sp. 1 0 0 0 0 0 0 0 0 0 0 1
Nichomachus minutus 0 0 0 0 1 1 0 1 - 1 1 1
Aopolonema princeps ? 0 0 0 1 0 0 0 0 0 1 1
Lattinova jacki 0 0 0 0 0 0 0 0 0 0 0 1
Acratheus ocellaris 0 0 0 0 1 1 1 1 - 1 1 1
Anapus americanus 1 1 0 0 0 0 0 0 1 1 1 1
Anapus kirshbaumi 1 1 0 0 0 1 0 0 1 1 1 1
Anapus rugicollis 1 1 0 0 0 1 0 0 1 1 1 1
Anapus dorsalis 1 0 0 0 0 1 0 0 1 1 1 1
Barbarosia punctulata 1 0 0 0 0 1 0 1 - 1 1 1
Chorosomella jakowleffi 0 0 0 0 1 1 0 1 - 1 1 0
Coridromius variegatus 0 0 0 0 1 1 0 1 - 1 1 0
Dasyscytus sordidus 0 0 0 0 0 1 0 1 - 1 1 1
Dicyphopsis sp. 0 0 0 0 0 0 1 1 - 1 1 1
Compositocoris senecionus 0 0 0 0 0 0 0 1 - 1 1 1
Dampierella schwartzi 1 0 0 0 0 0 0 1 - 1 1 1
Dimorphocoris gracilis 0 0 0 0 0 0 0 1 - 1 1 1
Dimorphocoris fuscus 0 0 0 0 0 0 0 1 - 1 1 1
Ectmetopterus micantulus 1 0 0 0 1 0 1 1 - 1 1 1
Euryopicoris nitidus 1 0 0 0 0 1 0 1 - 1 1 1
Goodeniaphila cassisi 1 0 0 0 0 0 0 1 - 1 1 1
Halticus apterus 0 0 0 0 0 1 0 1 - 1 1 1
Halticus intermedius 0 0 0 0 0 1 0 1 - 1 1 1
Halticus luteicollis 0 0 0 0 0 1 0 1 - 1 1 1
Labops burnmeisteri 1 1 0 1 0 0 0 0 1 1 1 1
Labops utahensis 1 1 0 1 0 0 0 0 1 1 1 1
Labops sahlbergii 1 1 0 1 0 0 0 0 1 1 1 1
Labops tumidiferons 1 1 0 1 0 0 0 0 1 1 1 1
Microtechnites altigena 0 0 0 0 0 1 0 1 - 1 1 1
Microtechnites bractatus 0 0 0 0 0 1 0 1 - 1 1 1
Myrmecophyes alboornatus 1 0 0 0 0 1 0 1 - 1 1 1
Myrmecophyes oregonensis 1 0 0 0 0 0 0 1 - 1 1 1
Namaquacapsus melanostethoides ? ? ? ? ? ? ? ? ? ? ? ?
Nanniella palustris 0 0 0 0 1 0 1 1 - 1 1 1
Orthocephalus arnoldii 1 0 1 0 1 0 0 1 - 1 1 1
Orthocephalus brevis 1 0 1 0 0 1 0 1 - 1 1 1
Pachytomella passerini ? 0 1 0 0 1 0 1 - 1 1 1
Piezocranum simulans 0 0 0 0 1 1 0 1 - 1 1 0
Plagiotylus maculatus 0 0 0 0 0 1 0 1 - 1 1 0
Schoenocoris flavomarginatus 1 0 0 0 0 1 0 1 - 1 1 1
Scirtetellus seminitens 1 0 0 1 0 0 0 0 1 1 1 1
Scirtetellus brevipennis 1 0 0 0 0 0 0 0 1 1 1 1
Strongylocoris leucocephalus 0 0 0 0 0 0 0 1 - 1 1 1

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

The Halticini of the world (Insecta: Heteroptera:

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

INTRODUCTION dark insects, often with saltatorial hindlegs and

ANIC, Australian National Insect Collection, CSIRO, TERMINOLOGY

KEY TO THE GENERA OF HALTICINI

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

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

Table 1. Host plant records for the Halticini

Genus Species Host plant Family Reference

Genus Species Host plant Family Reference

Remarks: This is one of two halticine genera endemic

Remarks: Compositocoris is very similar to Dimorpho-

Diagnosis: Halticus is characterized by a combination

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