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Home , Acalypta parvula, Acetropis, Adelphocoris seticornis, Agnocoris, Agnocoris reclairei, Amblytylus

A TAXONOMIC AND PHYLOGENETIC STUDY OF THE IMMATURE STAGES OF BRITISH (-)

by

George Cumming McGavin BSc (Hons), Edinburgh

A thesis submitted for the degree of Doctor of Philosophy of the University of London and ,for the Diploma of Imperial College

Department of Entomology British Museum (Natural History) London SW7 5BD

June, 1979 - i -

ABSTRACT

A taxonomic and phylogenetic study of the immature stages of British Miridae (Hemiptera-Heteroptera)

George Cumming McGavin

This thesis describes the Vth instar nymphs of the British Miridae, an heteropteran family of 206 . Keys to subfamilies, tribes and selected species are given, together with measurements for 24 body characters based on 10 individuals per species where possible. The dorsal abdominal gland openings are figured and the histology of the gland is examined. The morphology of the trichobothria occurring on the mesothoracic and metathoracic femora is investigated using scanning electron microscopy, and maps of their distribution in 116 species are drawn. The nymphal and adult patterns are found to be the same. The possible functions of trichobothria and their use in phylogenetic consideration of the group are examined. The structure of the pretarsus is found to be constant throughout the life cycle. One tribe, the , is investigated using numerical taxonomic techniques and the results are compared to those of conventional . Average linkage cluster- ing shows exact agreement with the latter. The interrelations of the genera of this tribe are examined using canonical variate analysis. CONTENTS page ABSTRACT INTRODUCTION 3 MATERIALS AND METHODS 14 Collection 14 Scanning electron microscopy 14 Slide mounts 15 Histology 16 Photography 16 Breeding 17 Numerical taxonomy 17 GENERAL MORPHOLOGY OF MIRID NYMPHS 19 Head 19 Rostrum 19 Antennae 20 Pronotum 20 Mesonotum and metanotum 21 Abdomen 21 Prothoracic leg 22 Mesothoracic leg 22 Metathoracic leg 23 Antennal growth 42 Campaniform sensillae 49 TRICHOBOTHRIA 55 60 Deraeocorinae 61 61 Dicyphinae 63 64 66 Discussion 68 DORSAL ABDOMINAL GLANDS 157 NUMERICAL TAXONOMY 195 Clustering analysis 196 Canonical variate analysis 202 page KEY TO THE SUBFAMILIES OF Vth INSTAR BRITISH MIRIDAE 214 Bryocorinae 217 Deraeocorinae 217 Phylinae 218 Dicyphinae 225 Orthotylinae 227 Mirinae 234 DESCRIPTIONS OF BRITISH MIRID NYMPHS 261 Bryocorinae 264 Deraeocorinae 266 Phylinae - 274 Dicyphinae 326 Orthotylinae 341 Mirinae 380 DISCUSSION 441

APPENDICES A Collectors 449 B Collecting sites 450 C Statistical data for the Stenodemini 452 D Altered county designations 459

ACKNOWLEDGEMENTS 460 BIBLIOGRAPHY 461 - iv -

List of figures page

1-4 SEMs - general features 25-32 5-8 SEMs - pretarsal structures 34-41 9-12 Antennal growth 43-46 13 Campaniform sensillae 50 14-18 SEMs - trichobothria 72-81 19 Femoral trichobothrial legends and maps 83-155 20-40 DAGO drawings 166-186 41-43 Micrographs and sections - DAGOs 188-193 44 Dendrogram 200 45-47 Canonical variate analysis 205-207 48 Ratios of antennal and body length in the Stenodemini 211 49-63 Key drawings 244-258 64 States of trichomae around a recessed bothrium 259 65 Body measurements 263 66 Deraeocorinae - head and pronotum 267 67 viridiflavus (Vth instar nymph) 276 68 arbustorum (Vth instar nymph) 309 69 pullis, C. saltitans and leucocephalus(Vth instar nymphs) 313 70,71 Dicyphinae - head and pronotum 327 & 334 72 Cyllecoris histrionicus (Vth instar nymph) 354 73 Dryophilocoris flavoquadrimaculatus 357 (Vth instar nymph) 74 rugulipennis (Vth instar nymph) 384 75 tripustulatus (Vth instar nymph) 387 76 pinastri (Vth instar nymph) 393 77 quadripunctatus (Vth instar nymph) 405 78 Megacoelum infusum (Vth instar nymph) 416 79 Collecting sites 451 - v -

List of colour plates page

1 planicornis (adult) 2 2 (Vth instar nymph) 13 3 Orthonotus rufifrons (adult) 18 4 Orthonotus rufifrons (IIIrd instar nymph) 24 5 salicellus (Vth instar nymph) 33 6 Dryophilocoris flavoquadrimaculatus (Vth instar nymph) 48 7 Alloeotomus gothicus (Vth instar nymph) 54 8 Megacoelum infusum (IVth instar nymph) 71 9 Megacoelum infusum (IVth instar nymph) 82 10 (Vth instar nymph) 156 11 Pilophorus cinnamopterus (Vth instar nymph) 165 12 ruber (IVth instar nymph) 187 13 Deraeocoris ruber (Vth instar nymph) 194 14 tiliae (IVth and Vth instar nymphs) 213 15 (Vth instar nymph) 216 16 lineolatus (Vth instar nymph) 243 17 pallicornis (Vth instar nymph) 260 18 (Vth instar nymph) 330 19 Dicyphus errans (Vth instar nymph) 331 - 2 -

Plate 1 Heterotoma planicornis (adult) lxLO 3

INTRODUCTION

Although the majority of systematics has been based on characters of the adult stages, the importance of the immatures has been recognised by a number of authors (Hayes,1931; Jordan,1951; Van Emden,1955 & 1957; Stammer, 1957 and Klausnitzer,1969). Characters of the immatures have been used to support the primary division of the Coleoptera, Diptera and and are as important • as the adult characters in the subdivision of the Odonata and the Neuroptera. Advances in the taxonomy and classifi- cation of the and have been made through studies of the nymphal* stages (Borner,1953; Borkhsenius, 1950). Van Emden (1955) gave examples of new families, tribes and genera and the transfer of species and species groups from one taxon to another, based upon the study of immature stages. Bibliographies of important works on the classification of the larvae and nymphs of various insect orders has been given by Van Emden (1955 & 1957), Stammer (1957) and Klausnitzer (1969).

In theory, characters of the adult and immature stages are of equal importance in taxonomy since both are derived from the same gene pool. A study of the immatures, however, offers a number of advantages. Comparative studies of immature forms are valuable in attempts to correlate onto- geny with phylogeny, sometimes affording better characters for examination than the adults.

Characters of immature will often define the same groups as will different characters in the adult insects. These are known as congruent groups. There are partial degrees of congruence and where characters of the immature stages define different groups; they are termed incongruent (Van Emden,1957). Some immature stages are of course secondarily adapted to their habitat, where this differs

The term nymph is used to mean the immature stage of hemi- metabolous insects - those with direct metamorphosis - as opposed to the term larva which refers to the immature stage of holometabolous insects - those with indirect or complete metamorphosis. - 4 - from that of the adults. In some aquatic orders of hemi- metabolous insects secondary adaptations are a result of differences in biology between the aquatic nymphs and terrestrial adults. Where a resting pupal stage occurs, the characters may also be unrelated. In practice, either the adult or the immature stage of an insect species may exhibit more accessible, less variable and more useful characters for classification, but in general the adult shows a greater degree of differentiation than the immature and is thus more suited to species identification. How- ever, the characters of immature stages are likely to be of greater importance for classification in the broader sense, such as the definition of higher categories (Van Emden,1955). The majority of literature shows that, by and large, the characters of immature stages can be used in the same way as those of the adult stages.

It has been stated that, in general, immature insects have a greater effect upon man and are more injurious than the adult stage, due to their presence in large numbers and for longer periods (Van Emden,1957), and their taxonomic study will therefore often be of great economic value. Nymphs and larvae of pest species are better targets for control measures, as in general their cuticle is softer and more vulnerable to various types of treatment. Immatures neither reproduce nor fly, which is beneficial in the field of biological control where parasites and predators have more chance of being effective. In general, control measures directed against immature stages are more likely to be quicker acting.

Knowledge of the taxonomy of larval and nymphal stages is also of use to ecologists who 'require such information in quantitative ecological research. When studying complex habitats and examining the food ecology and energetics of ecological systems, it is necessary to be able to determine the species of larvae and nymphs which are present. It may be that one or more closely related species appear to coexist, but when the immature stages are examined, differ- ences in their biology, food preferences, etc., may be found. The study of the ontogeny of characters in hemimetabolous insects, and the more complex changes of characters and character complexes in the development of holometabolous insects, will provide interesting insights into the gene- tic mechanisms controlling their inheritance and occurrence in the adult stages. Each cell of an insect species contains the same DNA, and Wigglesworth (1975) has shown that each cell must contain the metabolic systems needed for the formation of both larval and adult cuticle. The role of hormones is to determine which of these systems is effective. The systems involved in switching on sections of the DNA, allowing them to be expressed, and in switching sections off, have not been studied in great detail, although a certain amount of work in holometabolous insects has been done, mainly on Drosophila species (Lawrence & Morata,1972). In hemimetabolous insects, the expression of nymphal char- acters of the cuticle, specialised hairs and patterns, and the suppression of adult ones have been studied in prolixus (Wigglesworth,1975) and Oncopeltus fasciatus ( Lawrence,1966 & 1969). According to Wigglesworth (1972), the messenger RNA (mRNA) carrying coded instructions from the nucleus to the cytoplasm initiates a specific pathway of development by evoking the formation of particular enz- ymes, the cell undergoing a corresponding differentiation. He envisaged the cells of Rhodnius as carrying two families of mRNA, one maintained in action by the presence of juve- nile hormone, the other becoming activated by its absence. There will of course be other mRNAs which will be active in both nymphal and adult development. The study of immature stages of insects will facilitate this kind of work and lead to a better understanding of the developmental processes in the acquisition of adult characters. The surface morpho- logy and shape of cuticular sculpturing and hairs are indicators of the developmental condition of the test and, as the cuticle is secreted by a single layer of cells, it is a relatively easy system to analyse.

Taxonomic studies of the eggs of insects have been of great value in the classification of many groups, for example, Diptera (Carpenter & La Casse, 1955), Coccidae (Puchkova, (1955) and many others. This is especially so in the Het- - 6 - eroptera, (Heidemann,1911; Leston,1953; Southwood,1956a; Cobben,1968). Southwood (1956a) showed that the development, hatching and structure of the eggs of terrestrial Meter- optera supported the division of the Geocorisae into the Pentatomorpha and , and gave a good bibliography on previous works on eggs.

All parts of immature insects are potentially important in the process of classification. The central nervous system was used in a classification of closely allied larvae (Beier,1927), the tracheal system of larval diptera has been studied (Whitten,1959), and in immature Ephemeroptera, the tracheal system was used to show that the closed respirat- ory system of these orders arose from an open one (Calvert, 1929). Biochemical characters such as haemolymph proteins and enzymes may also be used. Brodie and Ryckman (1967) used agar gel diffusion and electrophoresis to study the ova antigens and haemolymph proteins in the Triatominae. They were able to identify all North American genera, species and subspecies, using nymphal haemolymph proteins. In addition 150 out of 180 fifth instar specimens were correc- tly sexed. Brezner and Enns (1958) used paper electrophoresis to distinguish between the larvae of Curculionidae and Scolytidae, and Norment et al.(1972) used acrylamide gel electrophoresis in taxonomic studies of Cicadellidae and Membracidae.

The scanning electron microscope has been recognised as being of use in the study of insect morphology and taxonomy, (Sakeld & Wilkes,1968; Taylor & Beaton,1970; Heywood 1971), especially where weakly sclerotised or fine structures needed to be resolved in detail. Its use in the taxonomy of larval Culicidae (Dah1,1972), mites (Sasa,1970), collembola (Lawrence & Massoud,1973), and many other taxa has been demonstrated. Hinton (1967) studied the larval spiracles of some Scaraboidea, and Raybould et al.(1971) found SEM studies of the larval cuticular patterns in African phoretic Simullidae of great value in species identification. Bacetti & Capra (1970) gave a brief review of the use of the SEM in systematic entomology, along with other references. - 7 -

The Hemiptera are divided into two suborders - the Heteroptera and the Homoptera - based on characters of the adult wing and rostrum. The former suborder contains, among others, assassin bugs'(Reduvidae), shield bugs (), capsid bugs (Miridae), bed bugs () and all the aquatic families, whilst the latter contains (Aphididae), coccids (Coccidae), hoppers (Cicadellidae), white flies (Aleurodidae) and psyllids (). Kirkaldy emphasised the importance of study- ing colour patterns and structures in hemipteran nymphs as a means of yielding generic and specific characters. He used the number of tarsal segments, the condition of the tegmina and the dorsal abdominal gland openings (DAGO) in the nymphal stages to construct a key to the families of Heteroptera (Kirkaldy, 1908a). There have since been several other keys to the families of Heteroptera based on nymphal characters. Hayes (1931) gave a good bibliography of the important works on the identification of nymphs, mentioning that at the time the only well worked group of terrestrial Heteroptera was the . In 1951, Jordan tabulated the number and position of DAGOs in nymphal Heteroptera and included the information along with colour and ecological characters in a key to families. The British heteropteran nymphs were dealt with by Leston & Scudder (1956). A key to the common families of Heteroptera was produced by Lawson (1959) and in 1961, Puchkov published: the characters of the nymphal stages of terrestrial Hemiptera in which subfamilies, tribes and genera were some- times distinguished on the basis of nymphal characters. He gave a key to selected species, which worked for IIIrd to Vth instars, based on colour, shape and type of pubesc- ence. He recognised the difficulty in determining species from Ist instar specimens, and Cobben (1978) stated that chaetotaxy of the Ist instar nymph is too inconsistent within larger taxonomic units to allow useful speculation to be made on its phylogenetic importance. It is, however, often possible to determine species from the Ist instar specimens where unique features are displayed. DeCoursey (1971) published a key to the families and subfamilies of North American heteropteran nymphs, and Herring & Ashlock (1971) gave a key to the nymphs of American families of - 8 - Heteroptera north of Mexico. These family and subfamily keys used the number and placement of DAGOs, the occurrence and distribution of trichobothria, the number of antennal segments, the shape and number of rostral segments as well as colour and biological data. Surprisingly little attention was paid to the characters of the pretarsus, although these had been shown to be useful (Dashman,1954; Goel & Schaefer,1970). Keys to determine the instar of heteropteran nymphs, based on the relative lengths of the mesothoracic and metathoracic wing pads, have been con- structed by Puchkov & Puchkova (1956) and Southwood (1956b).

Keys have been written for the nymphs of various families and subfamilies of Heteroptera. Slater (1951) produced a key to the nymphs of midwestern American , and the Russian Lygaeidae were dealt with by Puchkov (1958). Eyles (1963) studied five species of rhyparochromine Lygae- idae, and Scudder (1976) has also studied some nymphal Lygaeidae. The tribal classification of the Lygaeidae has been examined and modified, and nymphal characters were found to be of value in producing a balanced classification of the group, (Slater & Sweet,1961; Sweet,1967). Sweet & Slater (1961) have produced a key to 53 genera of nearctic Lygaeidae based on the Vth instar nymph. The nymphs of the were studied by Wheeler & Henry (1978) and pentatomid nymphs by Puchkov (1959a), DeCoursey & Esselbaugh (1962), McDonald (1963), DeCoursey & Allen (1968) and Oetting & Yonke (1971). Puchkov (1959b) studied the nymphs of and Arenocoreidae in the eastern part of the USSR, while Kumar (1966) dealt with some Australian coreids. An attempt to key the last nymphal instar of the Dutch species of was made by Cobben & Pillot (1960) using dorsal hair covering, femoral spines•and other characters. The authors were able to key out the Ist instar nymphs to sub- family, and accurate chaetotaxy was employed to identify some species. The nymphs of the British were classified by Sands (1957); the key to species was based on eggs, the Ist and the Vth instars and used mainly cont- inuous meristic data and colour. Southwood & Fes kes(1961) gave keys , descriptions and measurements for the more common species of British . Nymphs of the - 9 - () have been studied and described by Southwood & Scudder (1956 a)and Stusak (1974), the latter author giving a key to the Vth instar nymphs of the genus. Puchkov (1971) studied the immature stages of the genus Dictyla (Tingidae), giving a key to the IIIrd-Vth instar for 8 species, based on abdominal spines, pubescence, antennal segments and ecological data. The nymphs of Gerris in British Columbia were dealt with by Scudder & Jamieson (1972), who gave keys to determine all five instars based on colour pattern and size.

In general, the characters possessed by a Vth instar heteropteran nymph are present in the Ist instar, but the relative shape of different parts of the body, such as antennae, rostra, etc., is more similar between different taxa in the Ist instar than in later instars. In the Vth instar allometric growth of various parts changes relative size, whilst colour patterns can also change throughout ontogeny. There are many important characters, however, that appear to be conserved throughout the nymphal instars, such as the shape and position of the DAGO, the pretarsal structures and the trichobothrial patterns. The relative length of tricobothria, however, may decrease compared to the length of the body throughout development. Adult Heteroptera lack the DAGO, however, having instead func- tional metasternal glands, whose form and secretions have been studied and found to be of taxonomic value in various groups (Johansson & Braten, 1970; Staddon & Thorne,1974; Aldrich & Yonke,1975; Prestwich,1976; petting & Yonke,1978). This study of gland secretions is a relatively new field of research, using gas chromatography and spectrographic analysis, and has a lot to offer in systematic entomology.

There are several characters of the nymphal stages which also occur in the adults, such as the femoral tricobothria, the pretarsal structures and the number of rostral and antennal segments. The form of the genitalia in the ð and has been of great value in the classification of the Heteroptera (Lacaze-Duthiers,1852; Kullenberg,1941; Slater,1950; Dupuis,1951; Scudder, 1959; Dupuis,1963). These are only slightly developed in the later stages of - 10 - the Vth instar, to a degree to which they can be seen in cleared preparations. They are slightly sclerotised and of little practical value as taxonomic characters. Adults are also more suitable for techniques such as cytotaxonomy where rapid cell division is required. This has been used in the Miridae and Nabidae (Leston,1957)andgave useful insights into the higher classification of the former.

The Miridae are a large cosmopolitan family of Heteroptera comprising more than 5000 described species. In Britain the family is represented by approximately 206 species, unevenly distributed among 6 subfamilies, the Bryocorinae, Dicyphinae, Phylinae, Orthotylinae, Mirinae and Deraeocorinae. Nymphs of the Miridae can be distinguished from other heterop- teran nymphs by the presence of a single dorsal abdominal gland (usually located between the 3rd and 4th abdominal tergites), four-segmented antennae and rostra, and the presence of trichobothria on only the mesothoracic and meta- thoracic femora. Many workers have published on the classi- fication and phylogenetic relationships of the family, basing their findings solely on adult characters (Reuter, 1905 & 1910; Tullgren,1918; Knight,1923; Kullenberg,1944; Slater,1950; Carvalho,1952; Kerzhner,1962; Schuh,1974). These are more plentiful than those of the nymphs, which have therefore been neglected. The possession by the adults of some taxa of a pronotal collar, of use in separ- ating subfamilies and tribes (Southwood & Leston,1959), is not shared in the nymphs. The adults exhibit a more varied range of hair types, associated with the presence of developed wings (themselves of some taxonomic significance), three tarsal segments (as opposed to two in the nymphs), and developed genitalia as already discussed. Akingbohungbe, (1974) reviewed some nymphal characters of the Miridae, and gave a key to subfamilies, based mainly on pretarsal and DAGO characters. He recognised six types of DAGO and tabulated their occurrence in 53 genera of principally nearctic Miridae. In an earlier study (1973) he briefly des- cribed the nymphs of 56 Wisconsin species.

The majority of taxonomic work on the nymphs of the Miridae has been confined to small numbers of species in certain habitats, to pest species, or to species with other features of interest. Butler (1923) gave brief descriptions of a number of British mirid nymphs and added four more descrip- tions in 1924, giving information on colour, host etc. Brown (1924& 1925) described Heterocordylus genistae (Scopoli) and Chlamydatus evanescens (Boheman). The apple capsid, P]esiocoris rugicollis (Fallen) and the common green capsid, pabulinus (L) were described and measured by Austin (1931). Boselli (1932) gave drawings, descriptions and notes on the biology of the Hemiptera associated with in Sicily, which included two species of mirid. Jewett & Townsend (1947) described the biology of dolabrata(L) and nasutus (Kirschbaum). Lygus vosseleri Poppius, a pest of cultivated cotton in east and central Africa, was studied by Geering (1953), and in the same year Van Dinther studied the mirids associated with wild mulberry in . Puchkov and Puchkova (1956) included descriptions, drawings and keys to 16 species of mirid nymph as pests of agricultural crops in the USSR. Fourteen species of mirid nymph associated with the stinging nettle ( dioica L) were described and illustrated by Southwood and Scudder (1956b), who gave a key separating them on meristic data and colour. Five mirid species occurring on broom (Sarothamnus scoparius (L)) were dealt with by Waloff & Southwood (1960). The feeding habits of the nymphs and adults of the latter group of species were examined by Dempster (1964), using the precipitin test (Dempster,1960). Buczek (1959) studied nymphs of the tribe Stenodemini in Czechoslovakia and Cobben (1960) described the nymphs and ecology of Chlamydatus saltitans (Fallen) and C. evanescens on Sedum sp., and paludum (Sahlberg) on Carex sp. in the Netherlands. The immature stages and biology of •pygmaeus (Zetterstedt) and Cyrtorhinus caricis (Fallen), predacious on delphacid nymphs on Juncus, were studied by Rothschild (1963). Morris (1965) gave descriptions, drawings and measurements of the nymphs of Psallus ambiguus ( Fallen) on apple trees in Kent. The ecology of Poppius has been well documented in Scotland on oats (Stewart,1969), and in Poland on rye (Bilewicz-Pawinska,1969). The biology and rearing of the mirids found on cocoa was - 12 - discussed by Kumar and Ansari (1974). Wheeler et al. (1975' described the nymphal stages of Deraeocoris nebulosus (Uhler), a predator of pests on ornamentals, and gave notes on the biology of this species. Wheeler and Henry (1976) described and figured the nymphs and adults of 6 species of mirid associated with the ornamental honeylocust tree in Pennsylvania, and gave notes on their biology.

Because the existing keys and descriptions are fragmentary the aim of this present study is to examine nymphal characters in the Miridae and to construct a key to as many British species as possible. - 1 3 -

Plate 2 Heterotonia }Jlanicornis (Vth instar) x20 - 14 - MATERIALS AND METHODS

Collection Nymphs were obtained from a variety of sources: live collecting in the field, rearing (a few species only), and exiting collections. Air-dried material was unsuitable for scanning electron microscopy, but could be mounted on slides. Material stored in spirit was suitable both for slide mounting and for scanning electron microscopy, although for the latter technique, some cleaning was some- times necessary. Live material was collected from various habitats by sweeping vegetation and beating the branches of trees and shrubs over a beating tray. Material obtained in this way was either kept alive for rearing to later instars, for observation or colour photography, or preserved in 70% alcohol (for slide mounting and general use), or in Dubosq-brasil solution (for histology). As most preservation media tended to fade the natural colours of the nymphs, a record of the colours in live specimens was made at the time of collection.

Scanning electron microscopy (SEMI Specimens selected for SEM were critical-point dried, freeze dried or left untreated. Critical-point drying gave the best and most consistent results. Specimens were dehydrated through 70, 80, 95 and 100% ethanol (up to 2 hours in each), and then transferred to acetone for a short period before critical-point drying, using CO2 as the transitional medium. This technique has been described by Hearle et al.(1972) and Harris et al.(1972). Sometimes cleaning was required, and a number of techniques were tried, including soaking in degreasing agents, immersion in liquid nitrogen and ultrasonic cleaning. The last of these, which has been used in other orders of insects (especially Coleoptera), proved to be unsuitable, as the vigorous nature of the process removed many fine structures, such as the trichobothria. Cleaning by dropping into liquid nitrogen and soaking in Inhibsol were satis- factory in most cases.

Specimens were then fixed to stubs under a stereobinocular - 15 - microscope using colloidal silver (Silver Dag) or araldite epoxy adhesive. When araldite was used, the stubs were placed in a 45°C oven overnight to harden. It was sometimes necessary to prop up long structures, like legs, with small pieces of glass.

Stubs were coated in a Polaron SEM coating unit E5000, which was subsequently fitted with a peltier effect cooled stage to reduce heating of the specimens and improve coating (Polaron Equipment Ltd). The coating metals used were gold and gold-palladium, deposition being controlled at 40mA to give a layer at least 20nm thick. Stubs were also coated on an Edwards coating unit, model 12E6/1605, to give a 10-20nm thick gold layer (Edwards High Vacuum Ltd).

Stubs were examined on a Cambridge Stereoscan 600 at 7-10kV and on a Cambridge Stereoscan MkIIa at 14-18kV. Photo- graphic records were taken using Polaroid and Ilford HP4 film.

Slide mounts Due to the large numbers of specimens handled in this study, and the fragile nature of the material, slide mounting was chosen as the most convenient method of storage. Disadvantages of this method are that the colours are lost and that the process is relatively time consuming. The general shape of the nymph is also sometimes obscured by the dorso-ventral flattening. There are,however, many advantages and the use of the keys will often necessitate the mounting of specimens. Numerical measurements,which were extensively used in this study, were greatly speeded up by the use of slides. In the descriptions all measure- ments were obtained in this way, and although in some cases the values given are not 'real' due to the flattening effect mentioned above, all the material was treated in a standard manner and measurements are thus comparable. Slide mounting also greatly facilitates the storage and retrievability of specimens, as well as increasing the shelf life. A further advantage is that high-power work and drawing is made possible. - 16 - The method used was as follows. Spirit or air-dried material was boiled in 70% EtOH for a few minutes and then macerated in 10% KOH (potassium hydroxide solution) at 80-90°C. All heating was done using a Techne Dri-block heater DBI. When maceration was com- plete, the specimens were washed in 70% EtOH to remove the KOH. The specimens were then cleared in chloral phenol ( 1:1 w/w chloral hydrate/phenol) at 80°C. The specimens were allowed to cool before mounting in Berlese's fluid (Eastop & Van Emden,1972). The slides were placed in an oven at 38°C to dry for about two weeks, and were then sealed by ringing first with Euparal and then Murrayite (GBI Labs Ltd, Denton). The slides were stored in drawers in clear plastic envelopes.

Low-power work and slide measurements were performed on a Wild 145 stereomicroscope, fitted with a measuring eyepiece graticule. Low-power drawings were made with a camera lucida attachment. Study of histological sections, trichobothrial patterns and all high-power work was carried out using a Wild M20 equipped for phase contrast. High- power drawings were made with a drawing tube attachment.

Histology Material was fixed in Dubosq-brasil solution (Pantin,1964), dehydrated through 80 and 90% EtOH and isopropyl alcohol. Peterfi's Celloidin-paraffin embedding procedure was then followed (Pantin,1964). Longitudinal serial sections of Vth instar nymphs were cut at 6-10p on a Cambridge rocking microtome. The sections were then mounted and stained with haemalum and eosin, Heidenhain's azan, Masson's trichrome or Mallory's phosphotungstic acid haematoxylin (PTAH). Haemalum and eosin gave the most consistent staining and caused least disruption of the sections. Some adult material was also sectioned and here the chitin was softened in Diaphanol prior to embedding.

Photography Photographic records of dorsal abdominal gland openings (DAGOs) and the gland sacs were made on a Leitz compound microscope, using differential interference and phase - 17 - contrast techniques. Live nymphs were photographed using a Hasselblad camera and Agfachrome professional colour film, from which Cibachrome prints were made.

Breeding It was sometimes necessary to rear nymphs of early instars in to obtain later stages. Rearing phytophagous nymphs and adult was quite difficult, as they required healthy host . Predacious nymphs were much easier to rear as they merely required a constant supply of suitable prey. Most species of common aphids were quite satisfactory. The nymphs were kept in small transparent plastic cages, normally used for rearing (Blackman, 1971). A piece of plant with aphids was introduced daily. The transparent lid enabled low-power microscopical observation of the feeding habits of the nymphs. Some were reared to the adult stage to check the nymphal identifications.

Numerical taxonomy The programmes used are described in the relevant section.

All measurements given in the text are in mm. - 18 -

Plate Orthonotus rufifrons (adult 0 25 - 19 - GENERAL MORPHOLOGY OF MIRID NYMPHS

A good key to the nymphs of the families of British Heteroptera has been given by Leston and Scudder (1956) and a key to instars has been given by Southwood (1956a). Various other authors have produced keys and descriptions of certain Miridae (see Introduction). Akingbohungbe (1974) has given a key to the subfamilies of Miridae.

In Southwood's key to instars, he stated that the size of the adults wings does not depend on the size of the wing pads in the nymphal instars. In some of the species studied in the present work, however, this was not found to be the case. When there is sexual dimorphism in the state of development of the adult wings, it was found that nymphs with small wing pads gave rise to brachypterous adults and nymphs with large wing pads gave rise to macropterous adults. The key, however, still works for most of the species examined. This is discussed for the species involved in the chapter on descriptions.

The majority of this study involves Vth instar nymphs, since they are larger and show the same features as the earlier instars. They have also been more frequently collected and are better represented in the major collect- ions examined. The nymphs of Miridae differ greatly in colour, shape, size and habit, but a description of the shared features is given.

Head The head is approximately triangular and bears ecdysial suture lines, one connecting the eyes and the other extending posteriad from the midpoint of this line to the back of the head and continuing down the midline of the nota . The eyes may be either remote from, or touching, the pronotum.

Rostrum The rostrum is four segmented and varies considerably in length. Slender, possibly tactile, hairs occur along its length, and a cluster of sensory papillae occurs on the apex of the fourth segment. - 20 - Antennae Antennae are four segmented, the first segment usually being the smallest. The fourth segment is normally invested with various sensory hairs and sensillae (basiconic pegs etc.). These have not been investigated previously, or in the present study, but can be distin- guished from the other setae by using the SEM. The fourth segment is normally also flexible as a result of a special spiral thickening, which may involve only the medial portion of the segment, or may extend basally and apically as well. In the adult the whole of the fourth segment is constructed in this manner, as is part of the third segment. The exact function of this peculiar structure is not clear, but it could be that flexibility is needed in the sensory areas to avoid damage from contact with sur- rounding objects. Maceration in KOH often resulted in the thinner parts of the spiral thickening being dissolved, leaving the thicker parts with the sensory hairs in the form of a coil $outhwood,1953). The four antennal segments may be differentially thickened or lengthened. An extreme case occurs in Heterotoma planicornis (Pallas), where the first and second antennal segments are greatly enlarged. A pair of apodemes extend from the second segment into the first, where muscles are attached in a pinnate arrangement. At their other end, these muscles are attached to the cuticle at the base of the first segment. Specialised antennal fossae, such as occur in the second segment of the . adult d of thoracica (Fallen), are not present in the nymphal stages, these structures being obtained in the imaginal moult.

Pronotum The pronotum is generally trapezoidal; in some species the lateral margins can be approximately parallel. In the subfamily Dicyphinae there may be an anterior con- striction to form a 'pro-collar'. The pronotum has a well marked medial ecdysial line extending from the anterior to the posterior margin. Pronounced calli are present, representing the inner attachments of the dorso-ventral muscles. The microcuticular sculpturing present elsewhere on the body is absent in areas of muscle attachment, this being particularly evident on the femora. - 21 - Mesonotum and metanotum The wing pads are produced in the regions of the mesonotum and the metanotum. The state of development of these pads can be used to determine the instar of a nymph (Southwood,1956a). There is no meta- thoracic scent gland opening or ostiole, such as is found in the adult bug (Johansson & Braten,1970). There are two pairs of large spiracles situated in the thorax, one pair between the pronotum and the mesonotum, and the other pair between the mesonotum and the metanotum.

Abdomen There are eleven abdominal segments, the tenth and eleventh being much reduced in size. Segments II - VIII bear a pair of lateral spiracles whose openings are simple except in the Dicyphinae, where they are expanded to twice the diameter of their associated tracheae. This is also the case, but to a lesser extent, for the Halticocorini. The reason for this expansion is not known. The tracheae are spirally thickened with a blind tracheal sac branching off near each spiracle. In general, the dorsal surface of the abdomen is more densely pubescent than the ventral surface. Many types of seta are to be found on the dorsum, whilst those on the venter are of the simple type which taper apically. The dorsal abdominal gland opening (DAGO) is situated in the midline of the body between the third and fourth segments. The structure of both the opening and the gland can be seen in SEM micrographs and in histological serial sections. The surface structure varies among taxa and has been used in the taxonomy of the nymphs by Akingbohungbe (1974), although restricted to only optical microscopy. The cuticle immediately surrounding the DAGO may appear darker, due either to the increased concentration of cuticular sculpturing in this region, or to the presence of pigmen- tation.

The developing genitalia are visible in slide mounts of Vth instar nymphs. In most cases it is possible to sex all but the youngest Vth instars; it is not possible to sex nymphs of earlier instars as the soft structures are macerated. In some species there is development of an anal cone or tube, as in Deraeocoris ruber (L). The - 22 - abdomen may also have rows of spots laterally, or trans- verse dark or light stripes, or other patterns of pigment- ation.

Prothoracic leg The prothoracic leg, usually the smallest of the three, possesses a group of 3-5 sensory bristles at the proximal end of the coxa in the midline. There are also 4, presumably mechanoreceptive, bristles arising from large sockets where the trochanter joins the coxa. In slide mounted specimens a thin flexible area of cuticle is visible on the trochanter, which appears to divide this segment into two. The loss of a leg usually occurs by fracture at this point. The femur has no trichobothria or special structures. The extensor and flexor tibialis apodemes can be seen extending through the femur from the tibio-femoral articulation. The tibia bears a comb-like device at its distal end, used for cleaning the antennae and also to push prey (in the case of predacious species) off the end of the rostrum. The comb can also be used for cleaning the meso- and metathoracic legs. In general it is positioned ventro-laterally, and consists of a straight row of 12-20 blunt spines guarded at each end by a much stouter bristle, which is usually spirally fluted. The tarsus is two segmented, each approximately the same size, or with the second segment much longer than the first. The pretarsus is attached apically, and consists of a pair - of variously shaped and striated claws, joined together at their base by a region of flexible cuticle. The unguitractor plate (UTP) is situated at the base of the claws, to which it is joined by flexible cuticle. The UTP has 2-3 trans- verse rows, each of 7-9 scales (figs. 5-8) From the distal end of the UTP there arises a pair of parempodia, which may be hair-, strap-, flap- or lobe-like. A pulvillus, which varies considerably in size, arises from the ventral surface of each claw. A stout apodeme extends through the tibia from the UTP.

Mesothoracic leg The mesothoracic leg is similar to the prothoracic leg, but is usually larger and lacks a comb. The femur possesses trkhobothria, which will be discussed in a later chapter. - 23 - Metathoracic leg The metathoracic leg is similar to the mesothoracic one, but is generally the largest of the three. The femur may be expanded and modified for a saltatorial habit as in sp., Chlamydatus sp., sp., and some species of Phytocoris, the expansion being necessary to contain the extensor muscles. The femur bears trichobothria which are usually greater in number than those on the mesothoracic femora. In two species, Stenodema calcaratum (Fallen) and S. trispinosum Reuter, the femur bears 1-2 thickened spine-like project- ions (fig. 19:208,210 ),the function of which is unknown.

Various parts of the body of nymphs and adults have been discussed by previous workers. Dashman (1953) has reviewed the terminology of the pretarsus and considers the use of the UTP as a taxonomic character (1954) in the adult. Goel (1972a) also reviewed the structure of the UTP in the Heteroptera. Schuh (1975) has discussed the nature, distribution and taxonomic importance of trichobothria in the Miridae, and also (1976) the structure and taxonomic importance of the pretarsal structures. The mouth part structures and feeding strategies in the Heteroptera have been reviewed by Cobben (1978). Plate 4 Orthonotus rufifrons (IIIrd instar) x30 - 25 -

Figure 1

Scanning electron micrographs - general features a Blepharidopterus angulatus (x2000) - tip of adult rostrum showing chemosensory papillae (cp) and tactile hairs (h) on the labia. b Deraeocoris ruber (x2200) - tip of Vth instar nymphal rostrum showing papillae and hairs as in a. Note the cuticular sculpturing beneath. c Dicyphus paDicornis (x2100) - Vth instar nymph, abdominal spiracle. Note the thickening rings inside the opening. d Dicyphus pallicornis (x565) - Vth instar nymph, poly- gonal or cellular cuticular sculpturing on the pro- notum. This pattern is found notably in the Bryocorinae. It is probable that the boundaries delimit the product of one epidermal cell. e (x200) - Vth instar nymph, abdom- inal spiracles. f Harpocera thoracica (x500) - IIIrd instar nymph, dorsal abdominal hairs, showing unique clavate structure in this species. - 26-

a b

c - 27 -

Figure 2

Scanning electron micrographs - general features a Harpocera thoracica (x1000) - IVth instar nymph, dorsal surface of terminal abdominal segment showing clumped cuticular sculpturing and clavate hairs. b Harpocera thoracica (x2000) - IIIrd instar nymph, coronate hairs on antennal segment III. c Calocoris quadripunctatus (x2000) - IIIrd instar nymph, coronate hairs on pronotum. d HaBodapus montandoni (x510) - adult; spatulate or swollen ended hairs are unique to the three British members of the tribe and occur all over the body in adults and nymphs. In Hallodapus the inter- ommatidial setae are simple (s), whereas in Systellonotus they are spatulate. e Hallodapus rufescens (x500) - adult d, specialised metafemoral sculpturing peculiar to Hallodapus. f ater (x520) - Vth instar nymph, specialised flabellate setae occurring with simple setae in the dorsal surface of the mesonotum. These hairs are found only in Capsus and extend over most of the dorsal surface of the body. - 2 8 -

i fy y 11'1 " v ` 11 j ri. f- . 1., i*: i f / ''1,1", ITM7/1 - 29 - Figure 3

Scanning electron micrographs - general features a (x2000) - Vth instar nymph, flabellate hairs on pronotum. Note the longitudinal striations and terminal projections. b Deraeocoris ruber (x1160) - Vth instar nymph, apices of hairs from a lateral abdominal tuft. c sulcatus (x1900) - Vth instar nymph, area of femoral cuticle showing the normal cuticular sculptur- ing absent where a muscle attachment occurs. d Capsodes sulcatus (x1900) - Vth instar nymph, femoral cuticle with large spine showing the normal structure of mirid spines, i.e. having a seam down the length of the shaft as opposed to the continuous longitudinal fluting seen in specialised hairs such as trichobothria. e filicis (x480) - Vth instar nymph, pro- notal sculpturing showing the typical cellular or poly- gonal pattern of the Bryocorinae. f (x1050) - Vth instar nymph, dorsal abdominal gland opening, cuticle showing polygonal pattern. Note the petal-like arrangement of epidermal cells (as marked by the lines of cuticular spinules) around the bases of the setae. -30-

►j!!lNWW1110 "`kY"1411. agio:"CIVZI. .

4" `Z.1 •10.10511Alralk‘klabA, :.+s.." s0 41tiab 4 2i4 ■ il a4...... m.....4,...... -n . 6.*I- 4' N44\,1%X ItiVI ia M ses^.= ' 1. + ,-r, wilwb,„,ziktUili MZ-41k 411%14,1414:k 16,61

Z! a.~ eL°~~~ ~a1i - 31 - Figure 4

Scanning electron micrographs - general features

a decolor (x550) - Vth instar nymph, DAGO. b Alloeotomus gothicus (x580) - Vth instar nymph, DAGO. Note the sinuate anterior bar and trace of polygonal pattern in the sculpturing just anterior to the opening. c Deraeocoris ruber (x5500) - Vth instar nymph, campani7 form sensillae on the metathoracic trochanter. These sensillae are found on the dorsal and ventral surfaces of all mirid trochanters in a regular and constant arrangement. They also occur near the antennal sockets and on rostral segment II. In other insect groups they have been shown to be strain gauges, providing sensory information about stresses acting on the cuticle. They are sensitive to flexing in the long axis, the sense cell being attached to the under side of the thin dome (d) which is supported by thick side walls (s). d (x1100) - Vth instar nymph, comb at apex of prothoracic tibia. Although damaged,this comb shows the basic mirid pattern with thick guard spines at either end. e Harpocera thoracica (x1000) -Nth instar nymph, meso- thoracic tibio -tarsal joint. f Dicyphus rhododendri (x600) - pollen grains of Rhododendron sp. adhering (possibly by spider silk) to the tarsus of a Vth instar nymph. - 32- - 3 3 -

Plate 5 Psallus salicellus (Vth instar) x40 -34- Figure 5

Scanning electron micrographs - pretarsal structures

1 - claw, 2 - parempodium, 3 - pulvillus, 4 - ungui- tractor plate, 5 - guard setae

a pteridis : Bryocorinae (x1000) - meta- thoracic pretarsus showing large lobe-like parempodia and small pulvilli. b Alloeotomus gothicus : Deraeocorini (x2000) - meta- thoracic pretarsus viewed from the ventral surface, showing the unguitractor plate with three rows of scales ( each scale probably the product of a single epidermal cell). The strap-like parempodia, character- istic of the Deraeocorinae, can be seen. c Alloeotomus gothicus : Deraeocorini (x1130) - meso- thoracic pretarsus viewed from the front, showing both pulvilli and one claw. Pulvilli are small and adpressed, the claw appearing thick basally. d Lopus decolor : (x2150) - metathoracic pre- tarsus viewed from front, showing a small section of the striate claw on one side and the beginning of the massive rugulose pulvilli unique to this species. The hair-or bristle-like parempodium can just be seen. e Harpocera thoracica : Phylini (x1000) - mesothoracic pretarsus. Claw very curved apically, striate and with large pulvilli. The bristle-like parempodia arise from the apex of the unguitractor plate which has three rows` of scales. f Dicyphus stachydis : Dicyphinae (x2200) - metathoracic pretarsus showing the long, striate claw with basal pro- jection characteristic of Dicyphus. Parempodia are hair- like (part of pulvillus detached).

All specimens are Vth instar nymphs.

'`

- - - 36 - Figure 6

Scanning electron micrographs - pretarsal structures

1 - claw, 2 - parempodium, 3 - pulvillus, 4 - ungui- tractor plate, 5 - guard setae a Dicyphus pallicornis : Dicyphinm {x1800) - meta- thoracic pretarsus showing the entire straight, striate claw with basal projection, large pulvilli and hair- like parempodia. b Dicyphus pallicornis : Dicyphinae (x2050) - meso- thoracic pretarsus showing parempodia with striations. the surface structure of the pulvillus and the claw is very similar and it is possible that the•dicyphine pulvillus is not homologous to those of other mirids, being rather an extension of the claw as an adaptation to specialised substrates ( it is well known that the host plants of many dicyphines possess glandular or adhesive hairs). c Dicyphus pallicornis : Dicyphinae (x2000) - meso- thoracic pretarsus viewed from ventral surface showing unguitractor plate with two rows of scales as opposed to three in the Deraeocorinae, Phylinae and Mirinae. d Dicyphus pallicornis : Dicyphinae (x5000) - meso- thoracic pretarsus as above. e Dicyphus rhododendri : Dicyphinae (x2000) - prothoracic pretarsus showing different features from other Dicyphus spp. The claw is short and curved but has similar sur- face striations, basal projections and pulvilli. Parem- podia are striate. The small hole on the basal pro- jection has so far only been found in this species and its function is unknown. f Dicyphus rhododendri : Dicyphinae (x2000) - meta- thoracic pretarsus showing striate parempodia.

All specimens are Vth instar nymphs.

- 38 - Figure 7

Scanning electron micrographs - pretarsal structures

1 - claw, 2 - parempodium, 3 _ pulvillus, 4 - ungui- tractor plate, 5 - guard setae a Heterotoma planicornis : (x2000) - meso- thoracic pretarsus showing large, flap-like parem- podia and broad unguitractor plate with only two rows of scales characteristic of the Orthotylinae. b Calocoris quadripunctatus : (x1000) - meta- thoracic pretarsus showing flap-like parempodia, small pulvilli and curved claw. c : Mirini (x550) - metathora- cic pretarsus showing flap-like parempodia, small pulvilli and unguitractor plate with three rows of scales. d Megacoelum infusum : Mirini (x540) - metathoracic pretarsus showing typical mirine configuration. e Capsus ater : Mirini (x500) - metathoracic pretarsus. f Capsus ater : Mirini (x2000) - metathoracic pretarsus from ventral surface showing unguitractor plate. The papillate regions on either side of the unguitractor plate occur in many mirids, but the function is unknown.

All specimens are Vth instar nymphs. -]9-

a - 40 - Figure 8

Scanning electron micrographs - pretarsal structures

1 - claw, 2 - parempoditm'►, 3 - pulvillus, 4 - ungui- tractor plate, 5 - guard setae

a Capsodes saltatus : Horistini (x1000) - mesothoracic pretarsus viewed laterally, showing striate claw, small pulvillus and flap-like,striate parempodia. The ungui- tractor plate has three rows of scales. b Stenodema calcaratum : Stenodemini (x1000) - meso- thoracic pretarsus showing flap-like parempodia and unguitractor plate with three rows of scales. c Stenodema calcaratum : Stenodemini (x1000) - meso- thoracic pretarsus viewed laterally. d Notostira elongata : Stenodemini (x500) - metathoracic pretarsus. e : Stenodemini (x500) - meta- thoracic pretarsus from the ventral surface, showing unguitractor plate with three rows of scales, nine in the lateral rows and eight in the central row (c.f. Megacoelum infusum, eight and seven).

All specimens are Vth instar nymphs. 1 -

b

d

e - 42 - Antennal growth

The growth of the antennae is of interest. From obser- vation it appears that, although the entire grows at very nearly the rate of the total body length, its individual segments grow at differing rates, which may have biological significance. The antennal segments in all instars of four species, Leptopterna dolabrata (L), Calocoris norvegicus (Gmelin), (L) striatus (L), were measured (using a minimum of 3 individuals for most points, but only 2 for the Ist instar of ). The logs of the individual segment lengths and the total antennal lengths were plotted against the log of the total body length throughout the nymphal development of the four species (figs. 9-12).

In all cases, the fourth segment showed negative allometric growth, indicated on the graphs by the lower slope of the line. The second and third segments showed positive allometric growth. The regression lines were not com- puted as the correlation is obviously very high.

Scanning electron microscopy reveals that the surface of the fourth segment is invested with sensory structures, such as hairs and smaller pegs. If the number of sense organs is a determining factor for the size of any segment,. then surface area and not just length will be a more adequate measure. Assuming the segments to be cylindrical, it is possible to calculate the surface area of each by 2rirL (given the length and the radius). The results for 3 species are given below - the change in surface area from Ist to Vth instar for each segment is shown in the right hand column (the surface areas are in mm2) (table I). It can be seen that the fourth s✓gment changes relatively little in surface area, while the second increases many times. It is likely that this is a general feature in the Miridae, and although variation in the degree of change of different segments throughout the family may occur, the fourth segment may only increase 1-4 times in surface area from Ist to Vth instar. The process is continued in the adult, i.e. there is no sudden increase in surface area as -43 -

LOG PART • OF BODY•

0.8'

07

06- TOTAL 0.5 • O

04- -REF

03-

02-

01-

0 0•

- 01-

-02

-

cm-

- 05-

-06-

0 0.1 1 0.2 013 041 05 06 1 07 081 LOG I II III IV V BODY L Fig. 9 Leptopterna dolabrata. The total antennal length and the antennal segment lengths for all instars are plotted as log values against log of total body length. -44-

REF

LOG PART' OF BODY.

0.6

0.5

0.4

0.3

0.2

0.1-

0.

-0.1•

-0.2

-03

-0.4

-0.5

-0.6-

-0.7-

0.9 LOG BODY L 0 10.1 III 03111 I V • 0.6 0.7 as

Fig.10 Calocoris norvegicus. The total antennal length and the antennal segment lengths for all instars are plotted as log values against the log of the total body length.

- 45 -

■

LOG PART. REF BODY L . OF BODY

TOTAL ANT L. -8 O

0 1 .2 1 -3 -4 1 -5 .16 7 1 •8 I .9 1.0 LOG BODY L. I II III IV V Fig. 11 Stenodema laevigatum. The total antennal length and the antennal segment lengths for all instars are plotted as log values against the log of the total body length. - 46 -

LOG REF PART OF BODY

A

0.5 0.6 0.8 0 BODY L I II III IV. V

Fig. 12 Miris striatus. The total antennal length and the antennal segment lengths for all instars are plotted as log values against the log of the total body length.

- 47 - Table I - Surface area of antennal segments

Instar SA

Species Ant. Ist Vth Change segment in SA Leptopterna dolabrata 1 0.076 0.932 12.2x 2 0.080 1.127 14.0x 3 0.087 0.447 5.5x 4 0.117 0.243 2.1x Oncotylus viridiflavis 1 0.032 0.683 21.3x 2 0.030 0.700 23.3x 3 0.029 0.391 13.4x 4 0.051 0.174 3.4x Heterocordylus tibialis 1 0.026 0.157 6.0x 2 0.028 0.292 10.4x 3 0.026 0.124 4.8x 4 0.046 0.074 1.6x

far as can be observed. If the sensory structures are related to the surface area, it may be that the number of these structures remains constant throughout ontogeny, elongation of the other segments, particulary the second, serving only to push the sensory structures farther from the body. It is certain that there are different types of sensillae on the antennae and these may be located on different areas of the fourth segmenta The use of SEM will determine the numbers and types of sensillae present, and whether any changes of number or type distribution occur throughout ontogeny, but these aspects have not been investigated in the present work.

The difference in growth ratio of antennal segments has been recorded in Heteroptera (Matsuda, 1962) and specifically in Miridae (Matsuda, 1963). The differences are in agreement with those found in the present study. Plate 6 Dryophilocoris flavoquadrimaculatus (Vth instar) x15 - 49 - Campaniform sensillae

The neural regulation of movement has been studied by various authors, and Wigglesworth (1965) examined the coordination required by insects when walking and gave a brief review of the work in this field. Campaniform sensillae in Periplaneta americana have been described by Pringle (1938), who examined their structure, function and physiology. The sensillae were found to be present on the maxillary palps, the femora, the trochanters (in four groups), on the outside of the tibiae (proximally) and on each of the tarsi (one sensillum on each tarsal segment).

The sensillae are present in many insect orders, such as Diptera, Coleoptera, Orthoptera etc., and may be universal. The structure of campaniform sensillae appears to be remarkedly uniform (see Wigglesworth,1965, for review and illustration), consisting of a thickened circular or eliptical ridge of cuticle, in the middle of which is a dome of thinner cuticle. The top of the dome sometimes has a minute depressed area, which presumably connects underneath the dome to the nerve cell body or scolopale (fig. 4c ). In slide preparations the domed area appears as a pore in transmitted light.

Pringle (1938) has shown that eliptical sensillae react to flexing along their long axes, and circular ones to flexion in any direction. Campaniform sensillae are commonly found at limb joints, where stresses are set up in the cuticle by muscular action, or by the weight of the insect's body, but they have also been found in antennae, the gills of ephemeropteran nymphs, the halteres of Diptera etc. (for review see Wigglesworth,1965). Sensillae often occur in groups, all those of a particular group often having the same orientation. Each group acts as a unit, responding to the same type of shear force (Pringle,1938).

The proprioceptive function of campaniform sensillae has been well established. In the phasmid, Carausius, Von Buddenbrock (1921) showed that sensillae at the base of each leg controlled the walking reflex. Amputation of - 50 -

COXA

Qdsensory bristles

A2

campaniform P '' EMUR sensil Iae TROCHANTER 0.13

0.18

Fig. 13 Al, A2 mirificus Vth instar nymph B1, B2 Dicyphus rhododendri Vth instar nymph C Position of rostral segments during feeding. - 51 - limbs caused a new walking pattern to be adopted, but when the insertion of sticks allowed the amputated stumps to contact the ground, normal walking was resumed. Tactile hairs on the tarsi are therefore not important - the sensory structures involved are higher up.the leg. The weight of an insect is slung from the six legs, and weight borne on a leg will cause stress, thus stimulating an appropriately placed campaniform sensillam. In Pediculus humanis, the distal surface of each trochanter bears a group of 5 campaniform sensillae, and Wigglesworth (1941) concluded that they are well placed to detect strains in the limbs caused by resistance to their movements. The righting reflex in Periplaneta appears to be mediated by canpaniform sensillae and not tactile hairs (Pringle,1938).

In the present work, campaniform sensillae of uniform structure have been found on all trochanters, on the head and on the rostrum of mirid nymphs(fig.13). The form of the sensillae (fig. 4c ) agrees with descriptions given by other authors, and they are well placed to be of proprio- ceptive value. One sensillum occurs laterally on the cuticle of each side of the head where the antenna articu- lates, and is presumably responding to the strains placed on the cuticle as a function of the position of the antenna. A ring of 4 or 5 sensillae occurs around the midpoint of the second labial segment (fig. 13).

Cobben (1978) has examined the mouthparts and feeding strategies in the Heteroptera, and has given valuable bibliographic information on feeding studies in various taxa. The occurrence of campaniform sensillae is not mentioned by Cobben, who has specialised in the aquatic families. He does not mention that flexion of the rostral segments when feeding occurs in the Pentatomorpha, Miridae and Tingidae (largely phytophagous taxa). The stylet bundle passes through the first rostral segment and is held firm; the second and third segments are folded back from the bundle during feeding and the stylets are again firmly held by the fourth segment, which also holds the stylet bundle on the desired spot prior to penetration (fig. 13c ). Predatory taxa such as Nabidae, Anthocoridae - 52 - and the aquatic families of Heteroptera appear to hold their rostra rigid, while the stylet bundles are intro- duced into the body of the prey. It is likely that the campaniform sensillae occur in those groups which flex their rostra when feeding - however, more information is required to support this. The sensillae are orientated with their long axes parallel to the long axis of the rostral segment, and are thus able to respond to flatten- ing and bending of that segment (as opposed to sideways bending). The degree of bending and flattening in the long axis of the segment is a function of the penetration of the stylet bundle. In the nymphs of Nabidae and Anthocoridae (2 species of each) campaniform sensillae are located elsewhere; 2 in the proximal 3 of the second rostral segment, and 1 midway down the fourth rostral segment in the Nabidae, and 2 apparently circular sensillae in the distal 3 of the second rostral segment in Anthocoridae. The ring arrangement seen in the nymphs of Miridae is not present. This presumably reflects the feeding strategies of the groups. The physiological and morphological study of these sensillae should provide interesting information on the feeding mechanisms in Heteroptera.

The trochanters of mirid nymphs have 2 groups of campaniform sensillae with approximately 6 in each group. One group is on the morphological posterior surface, and the other is on the morphological anterior surface (i.e. anterior and posterior as it is oriented in life). The sensillae are exactly similar to those described in other insects and are in the someplace N'on Buddenbrock,1921; Pringle,1938) . The structure of the trochanter in mirid nymphs is in agreement with that described• briefly for adults by Goel (1972b) . It is composed of 2 podomeres, a small - proximal one articulating with the coxa, and a large distal one, providing articular surfaces for the femur. The two podomeres are not articulated with each other, and behave as a single functional unit as first described by Southwood (1953) in the genus Orthotylus. Goel considered the Miridae to be a specialised family with specialised trochanters. He does not, however, mention the presence - 53 - of campaniform sensillae.

The groups of sensillae mentioned above occur only on the distal podomere of the trochanter, and it is certain that they are involved in neural arcs and chains of reflexes, through which activities such as walking and the righting reflex are accomplished. The legs of some species of Miridae, in both nymphs and adults, especially Phytocoris sp., seem to be readily shed during collecting. In many cases the fracture occurred between the two trochanteral podomeres, the distal podomere being shed with the femur and tibia. This may be of biological significance, since when this occurs, the campaniform sensillae catering for that leg are also lost. The insect is better off losing a leg than being captured by a predator, but after one leg has been shed from one side, another loss on the same side would be a severe handicap. The loss of three legs from one side would be disastrous. Shedding the campaniform sensillae with the lost -leg may alter the neurological balance in some way and raise the threshold of leg shedding on the same side. Complex neurological reflexes are present in walking and there seems no benefit in shedding the distal trochanteral podo- mere unless it is advantageous to lose all possible sensory input from the shed leg. The thresholds for leg shedding in response to capture in the pro-, meso-, and meta-thoracic legs may differ as well, it being less easy for the insect to shed a hind leg than, for example, a middle leg. The coxae have a small group of sensory bristles distally, which come into contact with the distal podomere of the trochanter, presumably at some point during the walking cycle; their exact function is not known. The study of the trochanteral campaniform sensillae and their role in the walking reflex, and their possible r r use as leg loss indicators would increase our understanding of underlying neural processes. - 5 4 -

Plate 7 Alloeotomus gothicus (Vth instar) x20 - 55 - TRICHOBOTHRIA

The structure, distribution and taxonomic importance of tri- chobothria (specialised sensory hairs) in adult Miridae has been studied by Schuh (1975), covering 50 species (some identified only to genus), including the Isometopinae and , in addition to the British subfamilies. He reviewed the occurrence of trichobothria in the Arthropoda, and pointed out that their number and spatial arrangement had been shown to be important in several groups, including the Arachnida and Hemiptera. Ruckes (1960) discussed the diagnostic value of trichobothria in pentatomid taxonomy with respect to their use for the classification of lesser taxa. In the present study the Vth instar nymphs of 116 species of Brtish Miridae have been examined.

The function of trichobothria is not fully understood, al- though it is now fairly clear that they evolved from normal setae and are mechanoreceptive. It is unlikely that they fulfil the same role in all the families of Hemiptera in which they occur.

The innervation of trichobothria in other groups has been studied by G5rner (1965), Gnatzy and Schmidt (1971) and Edwards and Palka (1974), whilst Foelix and Chu Wang (1973) have shown that many trichobothria possess a tubular body, a feature common in the innervation of most mechanorecep- tors. In Gerris remigis () the trichobothria of the front legs are used in a highly specialised prey- capture mechanism - the ends of the trichobothria dip into the water and are used to determine the point of origin of the ripples on the water surface (Murphy, 1971; Lawry,1973). It is likely that other specialised functions for the trichobothria, using equally complex neural pathways,will be found in other taxa.

Trichobothria in Miridae are visible with the aid of a stereo- binocular microscope, and can be recognised by the fact that they are generally longer and *more slender than the normal femoral setae, and occur on the mesothoracic and metathoracic femora. In fresh slide mounts, slight - 56 - pressure on the cover slip will cause the trichobothria to wave about readily in the mountant, thus distinguishing them from the less mobile setae on the femora. The use of phase contrast greatly facilitates the examination of their structure.

The general trichobothrium is composed of a long, slender, longitudinally fluted shaft or trich, which tapers slightly at the point of insertion into the bothrium (basal structure). The trich also appears to be hollow, as revealed by SEM examination of broken trichs (fig. 15b). The shafts of the trichobothria are held at angles varying between 45° and 90° to the femoral cuticle. The bothrium may be recessed to varying degrees, flush with the surround- ing cuticle, or tuberculate, that is, raised up in a more or less domelike structure. Surrounding the bothrium in most species is a trichome, a region of cuticular sculp- turing. The usual form of trichome is a compact area of higher concentration of microspinules around the bothrium, sometimes with an increased concentration of spinules on the side of the bothrium to which the trich normally leans. Some trichomae may even appear slightly ovoid, their long axis parallell to the long axis of the femora. The individual spinules of the trichomae may be thicker and shorter than the normal spinules of the femoral cuticular sculpturing. In certain groups the femora are devoid of sculpturing and the trichomae are diffuse areas extending some distance from the bothria, and comprised of sparse aggregations of spinules. In these groups the trichomae of adjacent trichobothria may fuse.

The various components of the trichobothrial appear to be linked, suggesting a common genetic mechanism. For instance, a tuberculate bothrium is always associated with the lack of a compact trichome and possession of a relatively short trich, whereas a recessed bothrium is associated with a compact trichome and a relatively long trich. Tuberculate and recessed bothria never occur on the same femur, but the degree of recession and the concentration of spinules forming the trichomae may vary slightly. - 57 - In the Miridae, trichobothria are undoubtedly mechano- receptors, and are situated on the femora of the meso- thorax (mesofemora) and metathorax (metafemora). The Gerridae and the are the only other heteropteran families with trichobothria in these positions, although in the Gerridae, the trichs are situated on the coxae and trochanters as well, and the trichobothria lack trichomae (Schuh,1975). The trichobothria occur along the distal half of the femora, and may be directed slightly ventrad or dorsad, and sometimes laterad. Laterally occurring trichs are the least common and generally tend to be quite short. It is possible that some of the trichs may inf act be displaced or touched by the tibiae when the latter are flexed.

The trichobothria may serve a function during flight, the longer trichs being displaced by the main air current as well as receiving information about minor lateral air currents. Both micropterous and brachypterous species possess them, however, and there seems to be no correlation between the development of the trichobothria and the degree of flight activity. Moreover, the nymphs possess these structures in the same pattern in all instars, the structure of the trichobothria themselves also showing little ontogenic change. If the trichobothria were associated with flight, one might expect them to be developed at a later stage. Under experimental conditions the trichobothria of spiders, house crickets and thysanurans have been shown to detect air movements (Hoffman,1967; Corner & Andrews, 1969; Slifer & Sekhon,1970; Edwards & Palka,1974), although this is not necessarily the information that the organism receives.

In Miridae trichobothria appear to be necessary throughout the entire life cycle and could possibly be responsible for obtaining information about the topography of the sub- strate, the nature of air movements around the body, or perhaps some aspects of prey capture. In relation to the last hypothesis, certain trichobothria showing high stability of form and location throughout the family (nos.2 and 3 on the metafemora and no.2 on the mesofemora) - 58 - were measured accurately from drawings in 116 species, and their length was calculated as a ratio of the length of the femora to give an index of relative length (a high value of 15-20 representing very short trichobothria, and 3-6 representing long trichobothria). Certain obligate predatory species (Tytthus pygmaeus, Cyrtorhinus caricis, Carapyloneura virgula (HerrictrSchUffer) do have very long trichobothria, and certain obligate phytophagous species (Harpocera thoracica, Oncotylus viridiflavus (Goeze), Megaloceraea recticornis (Geoffroy in Fourcroy)) have very short trichobothria. There are, however, exceptions to this - Megacoelum infusum (Herrick-S:hdffer), thought to be largely predacious, has short trichobothria, and Monalocoris filicis (L), a completely phytophagous species, has relatively long trichobothria, although in this species the number of trichobothria is greatly reduced on both femora. Lack of information on species food preferences in their natural habitat has prevented the calculation of a correlation coefficient to test the hypothesis properly. Some species are omnivorous, but may show marked preferences in the wild, which may not be apparent in laboratory cult- ure. Further work on the feeding ecology of certain species is required.

Electrophysiological studies of the trichobothria would be difficult due to the small size of the experimental material, but would provide interesting information about these specialised setae. It is clear that quite complex neural organisation involving trichobothria is possible, as in Gerris remigis (Lawry,1973).

In agreement with Schuh (1975), it was found that there was no sexual dimorphism in the taxa examined and there was no difference in the number or spatial arrangement of trichobothria or of bothrial type between nymphs and adults of the same species. However, in relation to the length of the legs, it appears that the trichobothria are longest in the Ist instar and shortest in the adult. There is also a change in the pattern of cuticular sculpturing between the nymphs and adults in the same species, the nymphs possessing sculpturing on the entire femur, while - 59 - the adult loses the sculpturing except in the immediate area of the bothrium, thus giving the appearance of stronger trichomae. In no case did an adult gain cuticu- lar sculpturing where it was not present in the nymph.

Inthe trichobothrialnnps (fig.19), the right metathoracic femur is drawn from the dorsal surface* and the meso- thoracic femur is figured underneath, also as viewed from the dorsal surface. Schuh(1975) drew the left meso- and metathoracic femora as viewed from the ventral surface, although this does not alter the interpretation and compri- son of results. The majority of trichobothria on the meta- femur are ventrally placed, pointing behind the plane of the paper in the drawing. Dorsal trichobothria can be visualised as standing out from the paper in live specimens, and lateral ones as lying on the paper. The trichobothria are labelled dorsal, ventral or lateral (D,V or .L respec- tively). Tuberculate bothria are indicated by an open dot at the base of the trich; recessed bothria are represen- ted by a solid dot. An estimation of the bothrial condition for each species from optical microscopy and SEM is given below the mesofemoral maps. Trichomae are shown as a ring or circle around the bothria, representing the margins. No indication of the strength of the trichomae is given - but see Figure 64 and also the legends to the maps of the species descriptions. The lengths of trichs shown are accurate, and missing trichs are shown as dotted lines or omitted altogether. In some species certain setae appear very trichobothria-like, but are not sufficiently clear to be included in the trichobothrial assessment. These are designated by a question mark (?) and may represent regressed or degenerate trichobothria, or indeed incipient ones.

Schuh (1975) used location and lengths of trichs and type of bothrial development to designate homologies, and suggested a hypothetical primitive complement of 7-8 metafemoral trichobothria and 6 mesofemoral trichobothria.

*Dorsal surface as seen in a slide mount - this is in fact the morphologically posterior surface. - 60 - The same criteria have been employed in the present study, in homologising the trichobothria of the nymphal Miridae examined. There appears to be a fairly common pattern with- in the family shown typically by figure 19:26, 27, 162, 163. The general number of 7-8 trichobothria on the metafemur and 6 on the mesofemur is in agreement with Schuh's findings. Those British species showing ant mimicry did not, however, show the reduced trichobothrial numbers found by Schuh in non-British species. The homologies of the mesa- femur were easy to establish, the pattern being more constant than in the metafemur, where some variation was found. Trichobothria 1,2,3 and 7 on the metafemur were the most stable - 4,5 and 6 showed instability and were often augmented by supernumerary trichobothria in a lateral or ventro-lateral position, with very weak or absent trichomae.

The following section is a summary of the trichobothrial condition in the nymphal stages of 116 species of British mirid.

Bryocorinae The two members of this subfamily in Britain belong to the tribe Bryocorini and both are associated with pteredophytes. Bryocorini Taxa examined: Monalocoris filicis(Fallen), Bryocoris pteridis (Fallen) (fig. 19:1-5)

Contrary to the findings of Schuh (1975), the tricho- bothrial distribution and type does not conform to the hypothetical primitive pattern. The pattern found was unique - 3-4 metafemoral and 3 mesofemoral trichobothria. Trichomae were absent and the bothria were flush or slightly tuberculate. The British taxa dealt with do not appear closely related to those examined by Schuh, namely sp., sp., Sinervus sp., and Pachymerocerus pilosus Carvalho. They appear in fact to be more closely related to and of the Monaloniini, and some members of the Odoniellini. - 61 - There are also some similarities with the aberrant phyline, Harpocera thoracica, but the trichs in this species are very short. It is not known why Monalocoris filicis and Bryocoris pteridis should differ so radically from the other members of the Bryocorini mentioned above, but it is possible that their specialised niche may be responsible for the differences that occur.

Deraeocorinae Deraeocorini Taxa examined: Deraeocoris lutescens(Schilling), D. olivaceus(Fabricius), D. ruber(L), D. scutellaris(Fabricius), Alloeotomus gothicus(Fallen) (fig 19:6-15 and SEM fig. 4a-c)

Members of this tribe possess 6-7 trichobothria on the metafemur and 5-6 on the mesofemur. The trichomae were conspicuous and diffuse over a fairly large area of cuticle. Schuh (1975) examined only one member of this tribe and found a trichobothrial number of 6:6. The trichomae present were comprised of longer spinules than in the taxa examined here, and he described the bothria as slightly recessed. The latter are in fact flush, in agreement with the findings for the British members of the tribe. The species examined by Schuh, Deraeocoris ostentans(Stal), must now be considered as unusual in having 6 tricho- bothria on both the meta- and the mesofemur. The relation- ships of the Deraeocorini are uncertain,but they are more closely related to the than to other British tribes.

Phylinae The trichobothrial numbers for this subfamily are 5-8 on the metafemur and 3-7 on the mesofemur. Schuh included the Pilophorini in this subfamily, whereas in this work they are placed in the Orthotylinae. Many species in the Phylinae approach the primitive pattern and possess compact trichomae around recessed bothria, although some - 62 - lack trichomae. In agreement with the findings of Schuh, this subfamily appears most closely related to the Orthotylinae. Phylini Taxa examined: Lopus decolor(Fallen), Oncotylus viridiflavus, Hoplomachus thunbergi(Fallen), Tinicephalus hortulanus(Meyer-DUr), molliculus(Fallen), Amblytylus nasatus(Kirschbaum), paykulli (Fallen), Orthonotus rufifrons(Fallen), Harpocera thoracica, Tytthus pygmaeus, Brachyarthrum limitatum Fieber, (L), Plesiodema pinetellum(Zetterstedt), Psallus alnicola Douglas & Scott, Atractotomus magnicornis(Fallen), A. mall (Meyer-DUr), A. mirificus Woodroffe, Plagiognathus albipennis(Fallen), P. arbustorum(Fabricius), P. chrysanthemi (Wolff), Monosynamma bohemani(Fallen), (Reuter), verbasci(Meyer-Dtir ), Salicarus roseri (Herrich-SchUffer), Sthenarus rotermundi (Scholtz), Asciodema obsoletum (Fieber) (fig. 19:16-67 and SEM figs. 14d-f, 15a-d)

This tribe has 5-8 metafemoral and 3-7 mesofemoral tricho- bothria, usually with developed, compact trichomae and recessed bothria, although some members show the apparently linked characters of weak trichomae and less recessed bothria. The inclusion of the aberrant species Harpocera thoracica in this tribe gives the trichobothrial numbers their lower limit. This species has various unique features in both the nymph and the adult, and should perhaps not be placed in this tribe, but it is uncertain to which other tribe it is related.

Schuh has placed the genus Tytthus in the tribe Leucophoropterini. In the present study, Tytthus pygmaeus is included in the Phylini, but in agreement with Schuh's findings for the Leucophoropterini, it has a trichobothrial number of 7:6, strong trichomae and well recessed bothria. - 63 - Oncotylus viridiflavus is noteworthy in having one of the highest trichobothrial numbers in the tribe, but having trichs which are much reduced in length, a feature it shares with Harpocera thoracica. Hallodapini Taxa examined: Hallodapus rufescens(Burmeister), Systellonotus triguttatus(L) (fig. 19:68-71 and SEM fig. 15e, f )

This tribe has 7 metafemoral and 5-6 mesofemoral tricho- bothria, with well developed, compact trichomae and recessed bothria. Systellonotus triguttatus is believed to be an ant mimic in the nymphal and adult ? stages, but does not have the reduced trichobothrial number found by Schuh (1974 & 1975) in other ant mimicking taxa, although Hallodapus rufescens appears to present a similar con- dition to that found by Schuh for H. quadrimaculatus Schuh. Ant-mimicking taxa have other specialised features (reviewed by Schuh,1974) and the taxa examined in the present study have been shown to possess a unique hair type. The two species of Hallodapus appear to have specialised cuticular sculpturing on certain areas of the metafemora (fig. 2e ), but this does not seem to affect the trichobothrial characters.

Dicyphinae Taxa examined: Dicyphus constrictus(Boheman), D. epilobii Reuter, D. errans(Wolff), D. stachydis Reuter, D. rhododendri Dolling, D. pallicornis(Meyer-Dtir), D. annulatus (Wolff), D. globulifer(Fallen), (Herrich-Schaffer), caliginosus (fig. 19;72-91 and SEM fig. 16a-c)

The range of trichobothrial number was found to be 4-8 metafemoral and 3-6 mesofemoral, the arrangement of the trichobothria being in many cases very close to the primitive hypothetical pattern for the Miridae. The trichomae of this subfamily are like those of the -64- Deraeocorinae in that they are diffuse rather than compact around the bothria. They are unlike Deraeocorinae, how- ever, in that the bothria are tuberculate rather than flush, and the concentration of the trichomal spinules is less. The occurrence of fusion between the trichomae of certain adjacent trichobothria is unique in the Miridae so far examined (fig. 19:72,74,76,78,79,82,84) Fusion usually occurs on the metafemora between trichs 2 and 3. The cuticular sculpturing comprising the trichomae is not concentrated and extends quite far out from the bothria, thus as metafemoral trichs 2 and 3 are close, the trichomae appear to fuse.

Dicyphus rhododendri shows a very aberrant trichobothrial pattern and lacks trichomae (fig. 19:80 and 81). It is clearly not closely related to the other British members of the genus Dicyphus, a fact borne out by examination of the pretarsus (fig. 6 e, f) .

Campyloneura virgula has a high number (8:6) of long trichobothria, but the trichomae are very weak. It is interesting to note that this species is thought to be totally parthenogenetic, males being very rare. No nymphs of the genus Macrolophus from Britain were obtained, but a French member of the genus, M. caliginosus, was examined and found to have a primitive arrangement of trichs, weak trichomae and tuberculate bothria. Schuh (1975) suggested that this subfamily may have a closer relationship with the Bryocorini on the basis of the trichobothria and the pretarsus. This possible relationship is not supported by examination of the British Bryocorinae. The trichobothrial condition in the Dicyphinae supports its elevation to sub- family status.

Orthotylinae The trichobothrial number of this subfamily was found to range between 4 and 9 metafemoral and 3 and 7 mesofemoral. The trichomae, when present, were compact and the bothria usually recessed. - 65 - Pilophorini Taxa examined: Pilophorus cinnamopterus (Kirschbaum), P. perplexes Douglas & Scott (fig. 19:92-95)

Six metafemoral and 5 mesofemoral trichobothria with compact trichomae and recessed bothria were present. Halticocorini Taxa examined: Halticus saltator (Geoffroy), Strongylocoris luridus(Fallen), Pachytomella parallela (Meyer-DUr), (Hahn) (fig. 19:96-103 and SEM fig. 16d-f)

The trichobothrial number is 7-9 metafemoral and 3-6 mesofemoral, with strong, compact trichomae and well developed bothria. The trichobothria are generally quite long. Strongylocoris luridus was unusual in that trichomae were absent, the trichobothrial number was greatly reduced (4:3) and the bothria were tuberculate, characters shared with the Bryocorini. On the basis of the trichobothria, it would seem that Strongylocoris is not closely related to the other members of the tribe. It also possesses certain other unique features (see description). Orthotylini Taxa examined: chlorizans (Panzer), Cyllecoris histrionicus(L), Dryophilocoris flavoquadrimaculatus(DeGeer), flavomaculatus(Fabricius), Heterocordylus tibialis(Hahn), Heterotoma planicornis, Blepharidopterus angulatus(Fallen), Pachylops bicolor Douglas & Scott, Orthotylus flavosparsus(Sahlberg), 0. adenocarpi(Perris), 0. flavinervis(Kirschbaum), 0. virescens (Douglas & Scott), 0. concolor(Kirschbaum), 0. prasinus(Fallen), Pseudoloxops coccineus (Meyer-DUr), Cyrtorhinus caricis, Neomecomma bilineatus(Fallen), ambulans(Fallen), M. dispar(Boheman) (fig .19:104-141 and SEM fig. 17a-f)

This tribe has 4-8 metafemoral and 4-7 mesofemoral - 66 - trichobothria. Trichomae are usually present and compact and the bothria recessed. Cyllecoris histrionicus, Dryophilocoris flavoquadrimaculatus and Blepharidopterus angulatus are unusual in this tribe in having reduced trichobothrial numbers, no trichomae and tuberculate bothria. It is also interesting to note that the length of the trich in these species is reduced.

Mirinae In agreement with Schuh(1975), many taxa were found with trichobothrial numbers higher than the primitive hypo- thetical state. The range was 5-10 metafemoral and 4-8 mesofemoral trichobothria. The trichomae were usually weak, and the spinules comprising them quite short, as opposed to those of the Phylini, which were relatively long. In many cases the density of spinules in the tricho- mae was only slightly greater than on the surrounding femoral cuticle. Pithanini Taxa examined: (Sahlberg) I\th inst. nymph maerkeli(Herrich-Schaffer) (fig. 19: 142 and 143)

This tribe contains Myrmecoris gracilis, an ant-mimicking species, the nymphs of which are also ant-mimetic. Contrary to the findings of Schuh, the trichobothrial complement of this specialised taxon does not show reduction of trichobothrial numbers, having 7 metafemoral and 6 mesofemoral trichobothria with compact trichomae and recessed bothria. The other British member of this tribe, Pithanus maerkeli, has a trichobothrial complement of 8:6. Mirini Taxa examined: Wagner, L. rugulipennis Poppius, (Fabricius), rubricatus(Fallen), 0. viscicola (Futon), 0. kalmi(L), , L. contaminatus(Fallen), L. spinolai(Meyer-Diir), Camptozygum pinastri(Fallen), Plesiocoris rugicollis, nigritus(Fallen), P. palustris(Reuter), P. unifasciatus (Fabricius), -67 - Chara_ochilus gyllenhali(Fallen), rufipennis(Fallen), Miris striatus, Calocoris quadripunctatus(Villers), C. stysi Wagner, C. alpestris(Meyer-DUr), C. norvegicus, C. roseomaculatus(DeGeer), Adelphocoris lineolatus(Goeze), Megacoelum infusurn, binotatus(Fabricius), (Costa), (Fabricius), Capsus ater(L), tunicatus(Fabricius) (fig. 19:144-201 and SEM fig. 18a-c)

The trichobothrial number in this tribe was found to be variable, with many species possessing a trichobothrial number higher than the hypothetical primitive number. Trichomae in this tribe were never very strong and the bothria not as recessed as in the Phylini. Horistini Taxa examined: (L), C. sulcatus(Fieber) (fig. 19:202-205 and SEM fig. 18d)

The trichobothrial complement for this tribe was found to be 6-7 metafemoral and 6 mesofemoral trichs. The two members of the tribe examined had relatively long trichs and recessed or slightly recessed bothria. The trichomae of the taxa examined were compact. It is possible that Prepops insitivus(Say), the only taxon in this tribe examined by Schuh, and having short trichs and flush bothria with obsolete trichomae, is not typical. Stenodemini Taxa examined: gimmerthali(Flor), Stenodema calcaratum, S. trispinosum, S. holsatum (Fabricius), S. laevigatum, Notostira elongata(Geoffroy), Megaloceraea recticornis, ruficornis(Geoffroy), Teratocoris antennatus(Boheman), T. viridis Douglas & Scott, T. caricis(Kirkaldy), T. saundersi Douglas & Scott, Leptopterna dolabrata, L. ferrugata(Fallen) (fig .19:206-233 and SEM fig. 18e, f ) - 68 - The species of this tribe possess 5-9 metafemoral and 4-8 mesofemoral trichobothria, with weak to strong, compact trichomae and recessed bothria. Acetropis gimmerthali is unusual, however, in having a reduced trichobothrial number (5:4), no trichomae and flush bothria. Species of the genus Teratocoris had slightly strenger trichomae than the rest of the tribe, and bothria occasionally well recessed. Schuh considers this tribe to have flush bothria, but this appears to be the exception rather than the rule.

Discussion The trichobothria in the nymphal stages in many taxa of British Miridae are remarkably constant in pattern and number. However, their study is still valuable for iden- tification at tribal level. Where the patterns and numbers do differ, they can be used to identify genera and species. The classification suggested by the nymphal trichobothria agrees with that suggested by other characters, such as pretarsal structures. It is not yet possible to state which type of trichobothrium is the more primitive and which the more specialised. However, the femoral tricho- bothria are a uniquely derived character in the Miridae, supporting the monophyletic nature of the family (Schuh,1975). Adults possess the same trichobothrial pattern and type as their nymphs (Schuh,1975 and the present work).

Schaefer (1975) described the numbers and arrangements of abdominal trichobothria in nymphs and adults of the Tricophora (i.e. the superfamilies , , , and Pentatomoidea). He found an ontogenetic increase in trichobothrial number - certain trichobothria could be predicted to appear at a certain stage of devel- opment. Each abdominal segment had various clusters of trichobothria which were constant in position, but not in trichobothrial complement. He concluded that there were 'trichobothria-determining' fields on each segment of the tricophoran abdomen, and within each field, depending on the hormone balance, certain setae could become tricho- bothria. A comparable situation has been found in Oncopeltus fasciatus, where a cell which forms a nymphal bristle may be induced to form an adult hair under certain - 69 - conditions (Lawrence,1966). In contrast to these two cases, no change in number or arrangement throughout ontogeny has been found for trichobothria in the Miridae.

Schaefer (1975) also observed that the abdominal tricho- bothria did not change in length during ontogeny, and from this concluded that their sensory function remained the same. The meso- and metathoracic femoral trichobothria in Miridae are also sensory structures. They appear very long in Ist instars and do not grow very much in length throughout the life cycle (table II, and c.f. the slow growth of the fourth antennal segment, pp. 42-47) .

Table II - Ratio of trichobothrial length to femoral length

I II III IV V adult Pilophorus cinnamopterus metafemur trich no. 2 4.50 4.95 6.21 7.79 11.30 10.36 ft ff ff 3 3.07 4.76 6.15 7.50 9.82 9.52 mesofemur ff ff 2 4.07 4.27 5.34 6.99 7.80 7.58 Heterocordylus tibialis metafemur trich no. 2 5.71 7.63 7.96 8.43 8.36 9.78 If II ff 3 4.72 5.78 f.12 7.50 8.07 11.61 mesofemur ff ff 2 5.12 6.03 6.96 6.33 7.71 7.88 Leptopterna dolabrata metafemur trich no. 2 5.39 6.56 7.43 11.22 12.25 f} If ff 3 3.97 5.00 7.22 8.51 11.80 - mesofemur ft ff 2 4.16 5.05 5.52 8.04 9.36 - The table sets out the ratio of trichobothrial length to femur length for three trichobothria in all instars of three species (n=2 in each case).

In Trichophora Schaefer (1975 ), found that the longest abdominal trichobothria were associated with ground living taxa, and the short ones with. taxa living on plants. In the present study an attempt was made to relate trichoboth- rial length to feeding habits, but no simple correlation was found. Such a correlation would help to elucidate the trichobothrial function(s).

One function which has been suggested, but which seems most unlikely in Miridae, is that of receptors in courtship fondling (Schaefer,1975). In most cases where this type of - 70 - behaviour is displayed, there is some sexual dimorphism, one sex having the more active role in receiving signals. No sexual dimorphism has been seen in mirid trichobothria, which are also present in all instars.

Draslar (1973) studied the properties of abdominal trich- obothria in Pyrrhocoris apterus(L), and found 3 morphol- ogically and functionally distinct types in a cluster. The longest type was excited by a deflection in one direction and inhibited by deflection in the opposite direction. The other two were excited by deflection in any direction from the resting position. They also differed in type of adaptation and occurrence of spontaneous activity. In mirid nymphs, although the surface morphology of trichobothria on any femur appears the same, (except that their lengths vary), the subcuticular and neural structure may differ and it is possible that different trichobothria respond to different stimuli. Draslar gives no clues as to the function of the trichomae and none is given here, but they do not appear to steer the trichobothria as has been suggested by some authors. Some trichobothria have no trichomae, while others such as in the Dicyphinae and Deraeocorinae, have diffuse trichomae. In mirid nymphs trichomae are of constant structure on any leg, although there may be differences in the concentration of the spinules comprising them.

In order to determine the exact function of trichobothria in heteropteran nymphs and adults, there is much need for detailed SEM and TEM study and more electrophysiological and behavioural data. In Miridae, trichobothria occur only on the meso- and metathoracic femora, which suggests an identity of function throughout the family. However,in other families, they occur on• other parts of the body so that they may serve more than one function in the Heter- optera as a whole. - 71 -

Plate 8 Megacoelum infusum (IVth instar) x15 The nymph is drawing the antennal segments through the comb on the prothoracic tibiae. The body is held nearly vertical, supported by the meso- and meta-thoracic legs as the antennae are very long. - 72 - Figure 14

Scanning electron micrographs - trichobothria a Alloeotomus gothicus : Deraeocorini (x530) - meta- femoral trichobothria of Vth instar nymph. b Deraeocoris olivaceus : Deraeocorini (x2000) - meta- femoral trichobothrium of Vth instar nymph. c Deraeocoris ruber : Deraeocorini (x2150) - metafemor- al trichobothrium of Vth instar nymph. d Lopus, decolor : Phylini (x1000) - metafemoral tri- chobothrium of Vth instar nymph. e Lopus decolor : Phylini (x2000) - as above. Note longitudinal fluting on trich. f Chlamydatus wilkinsoni : Phylini (x1000) - meta- femoral trichobothrium (ventral) of Vth instar nymph. - 73- - 74 - Figure 15

Scanning electron micrographs - trichobothria a Chlamydatus wilkinsoni : Phylini (x5000) -metafemoral trichobothrium (ventral) of Vth instar nymph. Note how trich tapers on insertion into the bothrium. b Chlamydatus wilkinsoni : Phylini (x5000) - metafemoral trichobothrium (ventral) of Vth instar nymph. The trich has broken off at the base and a small black lumen can be seen. c Monosynamma bohemani : Phylini (x2100) - metafemoral trichobothrium of Vth instar nymph. d Monosynamma bohemani : Phylini (x5200) - metafemoral trichobothrium (ventral) of Vth instar nymph showing longitudinal fluting on the trich. e Hallodapus montandoni : Hallodapini (x1000) - meta- femoral trichobothrium of adult. The nymphs show a similar pattern. f Hallodapus rufescens : Hallodapini (x1000) meta- femoral trichobothrium of adult. The nymphs show a similar pattern. - 75 -

b - 76 - Figure 16

Scanning electron micrographs - trichobothria a Dicyphus stachydis : Dicyphinae (x530) - metafemor- al trichobothria of Vth instar nymph. b Dicyphus pallicornis : Dicyphinae (x1050) - meta- femoral trichobothrium of Vth instar nymph. c Dicyphus rhododendri : Dicyphinae (x2200) - meso- femoral trichobothrium of Vth instar nymph. d Halticus saltator : Halticocorini (x2000) - meta- femoral trichobothrium of Vth instar nymph. e Halticus saltator : Halticocorini (x2000) - meta- femoral trichobothrium of Vth instar nymph ( trich broken). f Halticus saltator : Halticocorini (x5000) - meso- femoral trichobothrium of Vth instar nymph (trich broken). v Q) - 78 - Figure 17

Scanning electron micrographs - trichobothria a : Orthotylini (x2150) - meta- femoral trichobothrium of Vth instar nymph. b Heterotoma planicornis : Orthotylini (x180) - meso- femoral trichobothria of Vth instar nymph. c Heterotoma planicornis : Orthotylini (x2300) - meta- femoral trichobothrium of Vth instar nymph. d Heterotoma planicornis : Orthotylini (x4600) - meso- femoral trichobothrium (dorsal) of Vth instar nymph. e ,Blepharidopterus angulatus : Orthotylini (x1150) - Metafemoral trichobothrium of Vth instar nymph. f Blepharidopterus angulatus : Orthotylini (x5150) - metafemoral trichobothrium of Vth instar nymph. - 79- - 80 - Figure 18

Scanning electron micrographs - trichobothria a Calocoris quadripunctatus : Mirini (x1100) - meso- femoral trichobothrium of Vth instar nymph. b Phytocoris sp. : Mirini (x2000) - mesofemoral tricho- bothrium of Vth instar nymph. c Capsus, ater : Mirini (x1150) - mesofemoral tricho- bothrium of Vth instar nymph. d Capsodes sulcatus : Horistini (x1900) - metafemoral trichobothrium of Vth instar nymph. e Teratocoris saundersi : Stenodemini (x500) - meta- femoral trichobothria of Vth instar nymph. f Teratocoris saundersi : Stenodemini (x5000) - meta- femoral trichobothrium of Vth instar nymph. -o

0 - 82 -

,i Plate 9 Megacoelum infusum (IVth instar) x15

Fig. 19:1-233 Femoral trichobothrial maps - legend

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Monalocoris filicis (?) meta 3 - f/t 5.66 5.10 R 1 ~ (6) " 3 - f/t 6.00 4.85 2 meso 3 - f/t 6.70 3 Bryocoris pteridis meta 4 - f/t 4.70 5.62 R 4 " meso 3 - f/t 7.06 5

Deraeocoris lutescens meta 7 +w f 6.04 5.83 a 6 " " meso 6 +w f 6.38 7 D. olivaceus meta 7 +md f 7.73 7.63 a13 8 1 meso 6 +md f 6.68 9 0 D. ruber r meta 7 +m f 10.00 8.53 a 10 meso 6 +m f 6.75 11 D. scutellaris meta 7 +m f 8.46 6.75 a 12 " meso 6 +m f - 13 Alloeotomus gothicus meta 6 +md f 8.80 7.26 a 14 It It meso 5 +md f 9.00 15

Lopus decolor meta 6 + sr 14.60 8.20 p 16 meso 5 + sr 8.60 17 Oncotylus viridiflavis meta 8 ++ sr 20.46 - R 18 " " =meso 7 ++ sr 10.77 19 Hoplomachus thunbergi• meta 6 +wc sr 6.84 7.50 R 20 " It meso 5 +wc sr 7.50 21 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Tinicephalus hortulanus meta 6 +c r 7.57 8.20 R 22 tr meso 5 +c r 8.72 23 meta 7 +w sr 8.00 7.60 R 24 It It meso 6 +w sr 6.00 25 meta 7 +c r 6.00 5.79 (3 26 meso 6 +w r 5.26 27 Macrotylus paykulli meta 6 + sr 11.25 7.82 R 28 tt It meso 5 +w sr 7.28 29 Orthonotus rufifrons meta 6 ++c r 6.00 5.27 R 30 It rr meso 6 +c r 4.33 31 Harpocera thoracica meta 5 - t 20.00 - R 32 It meso 3 - t 23.75 33 Phylus coryli meta 6 +w f 8.00 7.30 a(3 34 It meso 5 +w f 8.88 35 Tytthus pygmaeus meta 7 ++c r 5.04 4.71 a 36 tr meso 6 ++c r 3.95 37 Br achy arthrun 1imitatum meta 8 - f/t 10.63 7.66 ? 38

It It meso 7 - f/t 11.75 39 Plesiodema pinetellum meta 5 +w f/t 6.75 6.75 ? 40 n rt meso 4 +w f/t 5.85 41 Psallus alnicoiā meta 6 +c. sr 7.75 5.80 R?a 42 meso 5 +c sr 5.00 43 Atractotomus magnicornis meta 7 +c r 6.75 6.68 ? 44 tr It meso 6 +c r 4.47 45 A. mall meta 7 +c r 6.25 5.20 aR 46 It meso 6 +c r 4.16 47 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. A. mirificus meta 8 r 5.16 4.76 ? 48 meso 7 r 3.95 49 Plagiognathus albipennis meta 7 +c r 6.05 5.33 R 50 f n meso 6 +c r 4.52 51 P. arbustorum meta 7 +c sr 8.86 6.51 a3 52 meso 6 +c sr 7.14 53 P. chrysanthemi meta 7 +c r 6.04 6.00 aR? 54 meso 5 +c r 5.06 55 Monos namma bohemani meta 6 +c r 5.54 5.07 R 56 meso 5 +c r 5.15 57 Chlamydatus pullus meta 7 rr 6.00 4.71 R 58 ► TI meso 6 rr 3.43 59 meta 7 r 7.03 6.33 a13 60 meso 6 r 5.35 61 Salicarus roseri meta 7 sr 7.47 6.75 a 62 meso 6 sr 5.82 63 Sthenarus rotermundi meta 7 sr 11.50 12.36 R?a 64 tr meso 6 sr 6.86 65 Asciodema obsoletum meta 6 sr 6.30 5.78 R 66 meso 6 sr 7.62 67

Hallodapus rufescens meta 7 ++c r 7.25 6.13 R?a 68 n meso 6 ++c r 7.20 69 Systellonotus triguttatus meta 7 ++c r 9.47 6.92 aR 70 n i meso 5 ++c r 5.46 71

Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Dicyphus constrictus meta 6 +ed t 5.67 5.00 ap 72 I I► meso 5 +ed t 4.32 73 D. eFilobii meta 7 +ed t 8.53 9.14 p 74 meso 6 +ed t 7.25 75 D. -erransI t meta 6 +ed t • 5.08 5.77 ap 76 meso 5 +ed t 4.85 77 D. stachydis meta 6 +ed t 5.00 5.00 p?a 78 It meso 5 +ed t 4.55 79 D. rhododendri meta 3 - t 9.70 - a 80 IT meso 3 - t 7.10 81 D. pallicornis meta 6 +ed t 5.31 6.34 p 82 It meso 4 +ed t 4.96 83 D. annulatus meta 4 + t 6.41 7.07 p?a 84 meso 3 + t 6.41 85 D. globulifer meta 4 +w t 12.20 9.38 p 86 1 meso 3 +w t 12.80 87 loneura virgula meta 8 +w t 5.94 6.10 a 88 Campy1 ►1 meso 6 +w t 4.00 89 Macrolophus caliginosus* meta 7 +w t 6.86 6.07 ? 90 11 It meso 6 +w t 5.71 91

horus cinnamopterus meta 6 +c r 11.30 9.82 ap 92 Pilop1 II meso 5 +c r 7.80 93 P. perplexus meta 6 +c r 8.75 7.84 ap 94 II meso 5 +c r 8.00 95 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Halticus saltator meta 7 ++c rr - - 13? 96 ~— ~— meso 6 ++c rr 4.17 97 Strongylocoris luridus meta 4 - t - 12.54 P 98 meso 3 t 8.54 99 Pachytomella parallela meta 6 ++c rr 3.14 2.61 R 100 ' " meso 5 ++c rr 3.33 101 Orthocephalus saltator meta 9 +w r 6.60 6.22 in 102 It meso 6 +w r 5.50 103

Malacocoris chlorizans meta 8 + r 7.28 7.97 aI3 104 It meso 6 + r 5.20 105 Cyllecoris histrionicus meta 4 - t - - aR 106 11 " meso 4 - t - 107 Dryophilocoris meta 5 - t - 25.2 aR 108 flavoquadrimaculatus meso 4 - t 20.20 109 Globiceps flavomaculatus meta 6 + r 6.12 5.34 aR 110 rr rr meso 6 + r 5.00 111 Heterocordylus tibialis meta 6 +c r 8.36 8.36 aP 112 It rr meso 6 +c r 8.53 113 Heterotoma planicornis meta 7 + r 6.86 9.36 a13 114 It It meso 6 + r 4.44 115 Blepharidopterus angulatus meta 5 - st 9.09 11.11 aI3 116 'r It meso 5 - st 7.00 117 Pachylops bicolor meta 8 + r 7.25 7.80 R?a 118 rr meso 7 + r 4.70 119 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig.' trichs trich 2/ trich 3/ habit part femur f rfmur no. Orthotylus flavosparsus meta 7 + r 9.46 10.14 P 120 f► rr meso 6 + r 5.66 121 0. adenocarpi meta 7 + r 8.53 8.53 3 122 fr meso 6 + r 6.05 123 O. f l avinervi s meta 7 + r 8.92 8.93 (3 124 fr meso 7 + r 6.20 125 0. virescens meta 7 +mc r 8.76 8.76 a3 126 meso 6 +mc r 6.09 127 0. concolor meta 7 +mc r 8.17 11.17 3?a 128 meso 6 +mc r 4.81 129 0. prasinus meta 7 +mc r 7.93 7.93 R?a 130 ir meso 6 +mc r 10.50 131 Pseudoloxops coccineus meta 7 +w sr 5.62 6.42 ? 132 rr meso 6 +w sr 4.89 133 Cyrtorhinus caricis meta 7 ++c r 4.75 4.75 a 134 TY ĪĪ - meso 6 ++c r 4.32 135 Neomecomma bilineatus meta 7 + r 14.20 9.09 3 136 fr meso 6 + r 7.20 137 meta 8 ++c rr 6.27 5.19 aR 138 meso 5 ++c rr 4.40 139 M. dispar meta 7 +mc rr 4.82 4.80 a13 140 ir meso 6 +mc rr 3.70 141

Pithanus maerkeli meta 8 +c r 5.41 4.97 a3 142 meso 6 +c r 4.87 143 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Lygus maritimus meta 7 +w sr 9.63 12.35 p 144 II meso 5 +w sr 7.50 145 L. rugulipennis meta 7 +wc sr 8.14 - p 146 t tt meso 6 +wc sr 6.20 147 Liocoris tripustulatus meta 9 +w r 6.36 7.00 p 148 --t ī'-- It meso 6 +w r 6.07 149 Orthops rubricatus meta 9 +w sr 7.84 7.55 ? 150 It meso 6 +w sr 5.35 151 0. visicola meta 8 - f 9.16 10.00 p 152 meso 6 - f 12.00 153 0. kalmi meta 7 +w sr 10.00 10.00 R?a 154 meso 5 +w sr 7.17 155 Lygocoris pabulinus meta 8 +w sr 10.23 8.80 p 156 IT It meso 7 +w sr 8.14 157 L. cantaminatus meta 8 + r 12.60 11.58 p 158 I! It meso 6 + r 9.72 159 L. spinolai? meta 8 + r 7.40 9.52 p 160 meso 7 + r 5.93 161 Camptozygum pinastri meta 7 +w sr 9.40 7.42 p 162 It It meso 6 +w sr 5.65 163 Plesiocoris rugicollis meta 7 + sr 5.80 7.82 p 164 It IV meso 7 +w sr 6.60 165 Polymerus palustris meta 10 +w r 5.20 6.25 p 166 It it meso 7 +w r 5.68 167 P. nigritus meta 10 + r 6.05 6.81 p 168 If meso 6 + r 6.41 169 Figure 19 - legend (continued)

Species Femur No. of Trichomae .Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Polymerus unif asciatus II II meta 9 + r 8.57 7.50 P 170 meso 7 + r 5.40 171 Charagochilus gyllenhali meta 9 +c r 6.55 5.24 R?a 172 meso 7 +c r 4.63 173 Dichrooscytus rufipennis IT meta 8 - f 13.46 25.25 P 174 fi meso 6 - f 6.36 175 Miris striatus ------f t meta 9 - f 12.00 - aR 176 meso 8 - f 12.00 177 (Figs.176 and 177 were drawn to the same size to show the similarity between them) Calocoris quadripunctatus meta 7 - st - 13.36 R?a 178 " meso 6 - st 11.90 179 C. stysi meta 7 +w f 10.66 9.00 aR 180 ō meso 5 +w f 7.43 181 "i C. alpestris II meta 9 +w sr 10.74 11.15 R 182 meso 7 +w sr 13.00 183 C. norvegicus meta 10 + sr 11.44 9.86 R 184 " meso 6 + sr 7.53 185 C. roseomaculatus meta 10 + sr 12.54 13.80 P 186 " meso 6 + sr 12.11 187 Adelphocoris lineolatus meta 9/10 + f 11.30 11.76 R 188 ' meso 7 + f 7.61 189 Megacoelum infusum meta 8 - t 19.28 22.50 aR 190 " meso 6 - t 13.46 191 Stenotus binotatus meta 9 +w f/t 9.92 8.06 R 192 meso 6 +w f/t 6.92 193 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Miridius quadrivirgatus meta 10 - f 10.57 16.44 13 194 meso 7 f 7.20 195 Phytocoris tiliae meta 8 +w sr 10.44 9.40 a 196 " meso 7 +w sr 8.08 197 Capsus ater meta 8 +w sr 7.03 7.28 13 198 " meso 7 +w sr 4.17 199 meta 7 +w sr 11.20 12.95 R 200 meso 5 +w sr 8.81 201

Capsodes gothicus meta 7 +c sr 6.16 5.84 137a 202 meso 6 +c sr 4.25 203 C. sulcatus meta 6 + r 6.27 6.90 3 204 meso 6 + r 5.35 205

Acetropis gimmerthali meta 5 -/+w f - - R 206 ti r► meso 4 -/+w f - 207 Stenodema calcaratum meta 7 + r 15.88 27.00 R 208 meso 6 + r 6.80 209 S. trispinosum meta 6 + r 21.53 21.50 R 210 'r meso 5 + r 8.58 211 S. holsatum meta 7 + r 20.50 27.33 R 212 meso 7 + r 7.05 213 S. laevigatum meta 7 + r 17.25 30.66 13 214 tt meso 6 + r 8.00 215 Figure 19 - legend (continued)

Species Femur No. of Trichomae Bothria Ratio of Ratio of Feeding Fig. trichs trich 2/ trich 3/ habit part femur femur no. Notostira elongata meta 9 +w sr 19,73 14.88 R 216 meso 8 +w sr 7.90 217 Megaloceraea recticornis meta 7 +w sr 31.50 22.23 R 218 n meso 6 +w sr 11,66 219 meta 6 +w sr 19.20 17.45 R 220 meso 6 +w sr 8.14 221 Teratocoris antennatus meta 7 +mc rr 7.14 10.00 R 222 r, 1 meso 6 +mc rr 5.84 223 T. viridis meta 7 -1-mc r 8.29 9.33 13 224 meso 6 +mc r " 6.45 225 T. caricis meta 9 +c r 9.06 12.08 3 226 1 meso 7 +c r 6.64 227 T. saundersi meta 8 + r 8.58 9.12 R 228 N meso 7 + r 8.16 229 1 Leptopterna dolabrata meta 9 + sr 12.25 21.00 R 230 11 meso 8 + sr 9.36 231 L. ferrugata meta 9 + sr 15.05 21.33 R 232 It meso 8 + sr 9.22 233

Abbreviations: + = present, - = absent, w = weak, m = marked, c = compact, d = diffuse, e = extensive, ++ = strong, f = flush, f/t = flush/tuberculate, t = tuberculate, st = slightly tuberculate, sr = slightly recessed, r = recessed, rr = strongly recessed a = predacious, p = phytophagous, see also Figure 64. * = not a British taxon. Femur lengths are in mm.

- 93 - Figure 19

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V -156 -

Plate 10 Pilophorus perplexus (Vth instar) x20 - 157 - DORSAL ABDOMINAL GLANDS

Dorsal abdominal glands are found in the nymphs of most Heteroptera, but not in the adults. In the latter the meta- thoracic glands take over the secretory function, though not necessarily with the same secretion or the same function. The occurrence of dorsal abdominal glands in the heteropteran nymphs has been of great value in the classification of the group. Many authors have used the number and distribution of the gland openings (DAGOs), along with other characters in constructing keys to families and subfamilies of heteropteran nymphs (Gulde,1902; Kirkaldy,1908b; Leston & S cudder,1956 ; Lawson,1959; Puchkov,1959a & b and 1961; McDonald,1963; DeCoursey & Allen,1968; Herring & Ashlock, 1971; DeCoursey,1971; Akingbohungbe,1974). Cobben (1978) has comprehensively reviewed the nymphal abdominal glands and has constructed a table of their occurrence based on the literature and his own work. He has also mentioned some taxa in which the nymphal glands are retained in some way in the adult stage, although perhaps with an altered function.

The primitive number of abdominal glands in the Heteroptera is considered to be four. They arise as median invaginations of the intersegmental cuticle between tergites 3 and 4, 4 and 5, 5 and 6, 6 and 7. No modern taxa possess four glands although remnants of the fourth gland (between tergites 6 and 7) can be seen in some dipsocoromorphan and cimicomorphan taxa.

DeCoursey (1971) has given a subfamily key to the nymphs of Miridae, in which he used predominantly characters of the pretarsus. The nymphs of the subfamilies Orthotylinae and Mirinae do not display the convergent and divergent (respec- tively) parempodia that are seen in the adult, and were not separated in the key. Akingbohungbe (1974), however, used the form of the DAGO which lies between the 3rd and 4th abdominal tergites. He recognised 6 types of DAGO, and tabulated their occurrence in the nymphs of 53 principally nearctic mirid genera. His work'included the subfamilies Cylapinae and Hyaliodinae, which are not examined in the present work. The latter subfamily was erected by - 158 - Akingbohungbe (1974) to contain the Hyaliocies group). In the Orthotylinae genera examined, he found only two types of DAGO - those with simple openings without any tendency towards secondary doubling,and those with or without a sclerotised bar (anterior bar in present study). The Mirinae on the other hand were found to have openings that tended to be secondarily doubled or completely doubled (except in Dichrooscytus and , where the DAGO was absent).

In the present study the DAGOs of 130 species of Mirid were examined (figs. 20-40 ) and histological sections were made for certain species. The gland sac is single and directed anteriorly from the opening. It was found that the DAGOs of nymphs of British Miridae in the subfamilies Orthotylinae and Mirinae did fit the basic pattern found by Akingbohungbe (1974), the former having a single opening and the latter a double opening. However, certain species have lost the DAGO, namely Dichrooscytus rufipennis and D. valesianus, and in Strongylocoris leucocephalus the DAGO is very small or absent. There is also some variation in the shape of the orthotyline DAGO, some of which (Cyllecoris histrionicus and Mecomma ambulans) can appear very like the mirine form. It is therefore proposed that the unguitractor plate (UTP) of the pretarsus provides a much more reliable character for the separation of the nymphs of the two sub- families, since the mirine UTP has three rows of scales, while the orthotyline UTP has only two, the central row being much reduced.

In the Bryocorinae, previous workers have stated that the DAGO is occluded by a thin membrane, a character unique to the Miridae (Aryeetey & Kumar,1973; Akingbohungbe,1974). The DAGOs of Monalocoris filicis and Bryocoris pteridis were examined in the present study and were not found to be occluded by a membrane; this is in agreement with the findings of Carayon (1977), who studied the morphology and histology of the DAGs of the bryocorine genera Sahlbergella, Distantiella, Odoniella, Lycidocoris and Helopeltis among others. Carayon cnnsidered the bryocorine nymphal gland to be highly active. Aryeetey & Kumar (1973) could not find any definite function - 159 - for the glands in the tribe Monaloniini, although they found that the gland was bilobed and histologically complex. The surface morphology of the bryocorine DAGOs examined in the present study appears similar to that of some Nirini, but the cellular cuticular sculpturing of the pretarsus and other characters deny any real affinity between the two tribes. The type 6 described by Akingbohungbe (1974) (DAGO with sclerotised bar, but opening occluded or completely absent) does not occur, although the other 5 types described are of some descriptive value.

The morphology of DAGOs in the tribes examined is set out below:

Bryocorini: Opening appearing double with anterior bar sinuate. Cellular sculpturing extending to opening. (fig. 20:1,2) Clivenemini: The single member of this tribe Bothynotus pilosus (Boheman) was not examined. Deraeocorini: Opening strongly doubled and sclerotised. Anterior bar sinuate. No cellular sculpturing. (fig. 20:4-8) Phylini: Opening single or double. Anterior bar often strong and sinuate. No cellular sculpturing. (figs. 21:1-8; 22:1-8; 23:1-8; 24:1-8; 25:1-8) Hallodapini: Opening small, single, oval with anterior bar straight or slightly curved. No cellular sculp- turing. (fig. 26:3-5) Dicyphini: Opening small, single, often pale and slit-like. Anterior bar very weak (except Campyloneura virgula). No cellular sculpturing (except C. virgula). (figs. 27:3-8; 28:1-3) Pilophorini: Opening single with anterior bar strong, well sclerotised, curved and extending well beyond the margins of the opening. No cellular sculpturing. (fig. 28:4-7) Halticocorini: Opening single, oval with anterior bar strong, straight or curved and often raised or embossed (except Strongylocoris luridus and S. leucocephalus, opening very small or absent). No cellular sculpturing. (fig. 29:1-8) - 160 - Orthotylini: Opening single, oval with anterior bar weak. (Anterior bar slightly sinuate in Cyllecotis histrionicus and Mecomma ambulans). No cellular sculpturing. (figs. 30:1-8; 31:1-8; 32:1,2) Pithanini: Myrmecoris gracilis - opening single, small, kidney shaped. Anterior bar weak. No cellu- lar sculpturing. Pithanus maerkeli - open- ing strongly doubled. Anterior bar strongly sinuate. No cellular sculpturing. (fig.32:3,4) Mirini: Opening strongly doubled with anterior bar sinuate or strongly sinuate (except Dichrooscytus rufipennis and D. valesianus - opening absent). No cellular sculpturing.(f igs. 32:5-8; 33:1-8; 34:1-8; 35:2-7; 36:2-8:337.118) Horistini: Opening cbuble,strongly sclerotised (fig.38:4-6) Stenodemini: Similar to Mirini. (The opening of Teratocoris has a tendency to appear less strongly doubled) (figs. 38:7,8; 39:1-8; 40:1-4) The area of cuticle around the DAGO of mirid nymphs does not seem to be ridged or thickened in any way, although this has been described in alydid nymphs, there being a ridged evaporated area which is very thick and serves to protect the nymphs from penetration of the secretion (Aldrich,Yonke & Oetting,1972).

The structure of the glands in various taxa of heteropteran nymphs and adults has been examined (Gulde,1902; Brindley, 1930; Dupuis,1947 ; Carayon,1950; Remold,1962; Stein,1969; Aldrich,Yonke & Oetting,1972; AryeeLey & Kumar,1973; Oetting & Yonke,1978; Cobben,1978). Stein (1969) described the fine structure of the DAGs of nymphs of Dysdercus intermedius, and found that there were two types of cell involved in the construction of the gland - epithelial and secretory cells. Aldrich gt al. (1972) examined the DAGs of three nymphal alydid species and found the histology to be similar in all three. The gland sacs were lined with thin cuticle pene- trated by ducts which were Also lined with cuticle. The ducts passed through a layer of epithelial cells and then into the secretory cells, where they terminated in an"end apparatus "(= 'kopfchen' of Schumacher & Stein,1971) . The - 161 - end apparatus was surrounded by microvilli and acted as the collecting mechanism for the secretions. Aldrich et al. (1972) proposed that precursors of the secretions were produced in different parts of the secretory cell and reacted enzymatically near the end apparatus so that the toxic substances produced would then pass down the cuticle lined duct into the reservoir. An exactly similar structure was described by Schumacher and Stein (1971) for the meta- thoracic gland of the adults of Dysdercus intermedius) using electron microscopy, and for the metathoracic scent gland of the adults of Ilyocoris by Staddon and Thorne (1973). In this latter work, the authors described the ductule leading from the secretory cell to the lumen of the scent reservoir, and showed the porous structure of the end-sac in the secretory cell, through which the secretions pass. All the secretory cells described to date are very similar in con- struction. The small DAG in Hebrus has a few large secretory cells (not more than 20) (Cobben,1978). The cells have a large vesicle with coarse radial striations, concentrating to a narrow lumen at one end, from which the efferent ductule originates and discharges into the central Ieservoir. The efferent ductule in this case lacks an end apparatus, but is swollen. Cobben did not find a muscular coat around the gland in Hebrus, and only described one type of gland cell as opposed to two as seen in some other taxa.

In the present study, the single gland sac in mirid nymphs could be examined by preparation as for slide mounts (see Materials and Methods), and dissection of the sac was possible complete with DAGO and adherent cuticle. The cuticle of the gland sac is much thinner than the normal external cuticle (figs.42g;43a,lf). Using differential interference contrast microscopy, it was possible to see a number of small regular structures on the surfaces of the sacs. These have been described by Carayon (1977) for Odoniella rubra, and were termed secretory canals. These are the cuticle-lined ducts as they connect to the central reservoir and can be clearly seen in figures 41a & 42a. If, as in the glands described by Stein(1969), Schumacher & Stein (1971), Aldrich et al. (1972) and Cobben (1978), one efferent duct carries the secretion of one secretory cell, then the numbers of'secretory canals' - 162 - or ducts represents the number of secretory cells surround- ing the gland sac. In mirid nymphs the size of the DAGO is related to the size of the sac, large sacs having large DAGOs and vice versa. In taxa such as the Dicyphini where the gland is relatively small, the number of secretory canals, and thus cells, is low (10-20). The approximate number of secretory canals is given for 7 mirid species (figs.41a-f; 42a ). The present study reveals that Monalocoris filicis has a relatively large gland (approxi- mately 100 secretory canals) whilst Carayon (1977) figured more than 100 for Odoniella rubra. It is not certain to what extent the number of cells can be compared to activity or rate of secretion, but it is interesting to note that the smallest glands examined were those in the Dicyphini, a tribe whose members are restricted to plants that have glandular adhesive hairs and are poisonous (e.g. Digitalis), and may therefore not require elaborate predator defence mechanisms, if indeed this is their function. It seems certain that defence was the primitive function. An exception among the Dicyphini is D. rhododendri Dolling, which is predacious on Masonaphis lambersi (MacGillivray) (Dolling,1972). This species has a larger gland sac than the rest of the genus Dicyphus. In section it can be seen that the glands of mirid nymphs are not surrounded by a muscular coat and have an opening muscle attached to the posterior lip of the DAGO, which continues posteriad and attaches to the posterior margin of the fourth abdominal segment. The secretory cells of nymphs nearing ecdysis appear to degenerate, as described for alydid nymphs by Aldrich et al. (1972). Staddon and Thorne (1973) discussed the innervation of the muscles controlling the reservoir aperture in Ilyocoris cimicoides and found that the muscles opened the slit, closure being effected by the natural elasticity of the gland opening. It is likely that this is the mode of action in the gland muscle of mirid nymphs. The cells on the dorsal side of the gland are much smaller than those on the ventral side. How- ever, they are certainly all secretory in nature, judging by the ductules arising from them. In some species the ducts appear to be branched. The secretory cells do not appear to be overlaid by a layer of epithelial cells through which the secretory ducts pass. - 163 - The composition of hetero.teran secretion, both nymphal and adult, has received much attention since the advent of gas chromatography and mass spectroscopy. It is generally accepted that the secretions of the nymphal DAG differ from the metathoracic secretions of the adults. This is certainly the case in Miridae; aspiration of large numbers of nymphs of any species produces no detectable odour. However, the introduction of one adult to the pdoter gives off a strong odour, not always disagreeable. (The inhalation of the secretions from 3 nymphs and one adult of Pentatoma rufipes (L) caused a sore throat, headache and nausea, which lasted for several hours . It is not certain whether the nymphs or the adult were responsible.) The difference in adult and nymphal secretions was noted by Aldrich and Yonke (1975) in coreids, although the nymphal and adult secretions of the cydnid Scaptocoris have both been shown to have fungicidal action against Fusarium spp.(Roth,1961; Timonin,1961). In the coreid spp.studied by Aldrich and Yonke (1975), the adult secretions were found to be composed of differing volatile aliphatic acids, aldehydes, alcohols and esters, variations in which were found to support a morphologically based classification. The nymphal secretions, however, were more uniform, being composed of an aqueous phase,and an organic phase containing trans-2-hexenal and trans-4-oxo-2- hexenal. Calam and Youdeowei (1968) reported the presence of n-dodecane, n-tridecane, n-pentadecane, hexenal, hex-2- en-1-al, 4-oxohex-2-en-1-al, oct-2-en-1-al and 4-oxooct-2- en-1-al from the posterior scent gland of the Vth instar of Dysdercus intermedius. Whole secretions and preparations of hexenal and trans-hex-2-en-1-al caused the dispersal of aggregations of bugs, and the function of an alerting phero- mone was ascribed to the secretion of the posterior gland.

Prestwich (1976) studied 8 species of east African Heterop- tera (4 pentatomids, 4 coreids) and corroborated the find- ings of Aldrich and Yonke (1975) in finding trans-2-hexenal and 4-oxo-trans-2-hexenal in the nymphal secretions. The immature coreids produce highly unsaturated aldehydes and ketones, unlike their respective adults which synthesise their secretions from acetone. The nymphal secretions exuded on the dorsal surface of the abdomen have low cuticle-penetrating - 164 - power, whereas the complex adult secretions with high cuticle-penetrating power are sprayed from the metathoracic glands (Prestwich, 1976).

Suggested functions for nymphal heteropteran secretions include alerting or alarm , defence or for aggregat- ion , as in 08 saldid Aepophilus (Baudin,1955). It may be that the DAGs%involved in the elimination of toxic substances from the food material ingested by some species. Waterhouse, Forss and Hackman (1961) remarked that the secretions of certain pentatomids and coreids contained compounds that occurred in the plants on which they fed. The sequestering of cardiac glycosides from certain Asclepiaeaceae by the monarch butterfly,Danaus plexippus, is a well documented case of an insect taking toxic compounds from plants, concentrat- ing them and using them as a defence mechanism (Erlich & Raven,1964). Aldrich and Yonke (1975) suggested that the distinctive trans-2-hexenal and trans-4-oxo-hexenal in the secretion of the nymphs of many taxa may have evolved via the selection by to mimic formicid alarm . Since their usual defence is to drop to the ground, they face a risk from ground and thus the production of ant alarm pheromones may be strongly selected. Cobben (1978) interpreted the existence of fully developed glands in the nymphs of Corixidae as being evidence that the secretions are of value in an aquatic mode of life. A defensive role for the secretions of the DAG in nymphal Miridae was not found by Aryeetey and Kumar (1973) in cocoa-feeding species, or by Remold (1962) in ten other species. The exact function of the DAG secretions in mirid nymphs is uncertain and awaits further investigation.

- 165 -

Plate 11 Pilophorus cinnamopterus (Vth instar) x25 - 166 - Figure 20

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BRYOCORINAE. Deraeocorin1. 1. 1ionalocor1s filicls x250. !4. Deraeocoris lutescens x250. 2. Bryocorls pterldls x250. 5. Deraeocorls olivaceus x250. DERAEOCORINAE. 6. Deraeocorls ruber x250. C11v1nemin1. 7. Deraeocorls scute11ar1s x125. 3. Eothynotus pilosus. N.S. 8. Alloeotonxts gothicus x250. - 167 - Figure 21

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PHTLINAE. 5. Conostethus roseas x125. Phylin1. 6.Hoplomachus thunbergi x125. I. Lopus decolor x250. 7.Tinicephalus hortulanus x250. 2. Oncotylus viriditlavis x250. 8. 1 galocoleus molliculus x250. 3. Conostethus brevis N.S. 4.Coristethus gr1seus N.S. - 168 - Figure 22

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1.Megalocoleus pilosus x125. 5.Macrotylus solltarlus x125. 2. Amblytylus brevicollis N.S. 6.Macrotylus paykulli x250. 3.Amblytylus delicates x125. 7.Orthonotuj rutifrons x250. 4. Amblytylus nasutu3 x250. 8.Harpocera thoracica x250.

- 16 9 - Figure 23

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1.Tytthus geminus N.S. 5.Phylus melanocephalus x250. 2. Tytttius pygnaeus 050 6.Phylus pa111ceps N.S. 3.Brachyarthrum limitatum x125. 7.Pleslodema pinetellum x250. 4.Phylus coryll x250. 8.Psallus ambiguus *

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1.Psallus alnicola x250. 5.Plaglognathus arbustorum x250. 2.Atractotomus magnlcornls x250. 5. Plaglognathus chrysantheml x250. 3.Atractotomus mall x250. 7.Plaglognathus vitellinus N.S. 4.Plagiognathus albipennls x250. 8.Ohlaanydatus pulicarius N.S.

- 171 - Figure 25

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1. Chlaa4Ydatus pullus..25o 5. tbnosynamma bohemani x125. 2. Chlarnrdatus saltftans x125. 6. Campyloama verbascf x250. 3. Chlamydatus wilkfnsoni N.S. 7. Salfcaras roseri x250. li. Chlamydatus evanescens N.S. 8. Sthenarus rotersmnd1 x250. - 172 - Figure 26

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1. Asciodems fiebert x250. 5. Systellonotus triauttatus x250. 2. Asciodema obsoletum x250. Hallodapini. 3.Hallodapus montandon1 x125. 1,. Hallodapus rsfescens x125. — 173 — Figure 27

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DICTPHINAE. I. tacrolophus nubills N.S. 5. Dlcyphus errans x250. 2.Macrolophus rubs N.S. 6. DIcyphus stachydls x250. 3.DIcyphus constrictus x250. 7.Dicyphus palllcornis x250. 4. Dicyphus epilobil x2500 8. Dlcyphus annulatus x250.

- 174 - Figure 28

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1.Dlcyphus globulifer x250. ORTHOTTLINAE. 2. Dlcyphus rhododendr1 x250. Pllophorinl. 3.Campyloneura virgula x250. 4. Pllophorus cinnamopterus x2500 5.Pllophorus clavatus x1250 6. N.S. 7.Pllophorus perplexus x250. - 175 -

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Halticocorini. 5.Strongylocoris leucocephalus 1. x125. (very small opening. ) 2.Halticus N.S. 6.Strongylocoris luridus x250. 3.Halticus saltator N.S. 7.Pachytomella parallele x125. 4.Halticus luteicollls N.S. 8.Orthocephalus saltator x250.

- 176 - Figure 30

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1. fialacocorls chlorizans x250. 5. Globlceps flavoraculatus x250. 2. Ffeberocapsus flaveolus x125. 6. Globlceps cruclatus x125. 3. Dryophflocoris flavaluadrfmaculatus x250. 7. Globlceps woodroffel x125. 4. Cyllecorts histrionfcus x250. 8. Heterocordylus genlstae x125. — 177 — Figure 31 1 • i

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1.Heterocordylus tibialis x250. Orthotylus omitted except, Orthotylus flavinervis x2500 2.Heterotonia planicornis x250. 5. Pseudoloxops coccineus x250. 3.Blepharidopterus angulatus x250. 6. cyrtorhinus caricis x125. 4.Pachylops bicolor x250. 7. 8.Neonecoama bilineatus x250.

- 178 - Figure 32

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Marini. I, Mecca= anbulans x125. 5.Lygus maritiaus x125. 2.Mecomma dispar x250. 6.? x250. MIRINAE. 7.Lygus punctatus x125. Pithanini. 8. x125. 3.Myrmecoris gracilis (4th.) x125. 4.Plthanus maerkell x250.

- 179 -

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1. Lygus rugulipennis x250. 5, Orthops rubricatus (4th) x250. 2. Liocorls tripustulatus x250. 6. Orthops viscicola x250. 3. Orthops atoaarius N.S. 7. Orthops campestris x125. 4. Orthops cervinus x125. 8. Orthops kalmi x250. - 180 - Figure 34

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1. Lygocoris pabulinus x250. 5. Lygocoris lucorum N.S. 2. Lygocoris contaminates x250. 6. Lygocoris spinolal x250. 3. Lygocoris popull N.S. 7. reclairel N.S. 14. Lygocoris viridis x125. 8. Camptozygum pinastri x250. - 181 - Figure 35

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1.Zygicus nigriceps N.S. 5.Polymerus vulneratus N.S. 2.Plesiocoria ruglcollis x250. 6.Polymerus nigritus x250. 3.Polymerus palustris x250. 7.Charagochilus gyllenhali x250. G. Polymerus unifasciatus x125. 8.Dichrooscytus rufipennis, (no visible opening.) - 182 - Figure 36

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1.Dlchrooscytus valeslanus. 5. Calocoris sexguttatus x250. (no visible opening.) 6. x250. 2.Miris striatus x125.(slide.) 7. Calocoris norvegicus x250. 3.Calocoris tulvomaculatus x125. 8. x250. 4.Calocoris quadripunctatus x250. - 183 - Figure 37

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1. Adelphocorls Ilneolatus 1250. 5. tiegaeoelum lntusum (771 ) x250. 2. Adelphocorls setlcornls (4th) X125. 6. Stenotus blnotatus 1250. 3. Adelphocorls ttclnensls x125. 7. tilrldlus quadrlvlrgatus N,S. PhYtOcorls omitted except. 80 Phytoeorls longlpennls x250. - 184 - Figure 38

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1.Capsus ater x2500 5.Capsodes gothlcus x250. 2.Capus wagnerl N.S. 6.Capsodes sulcatus x125. 3.Pantllius tunicatus (4th) x125. Stenodeminl. Horlstlnl. 7. Acetropls gimmerthall x250. 4. (4th) x125. 8.Stenodeva calcaratum x250. — 185 —

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1.Stenodema trispinosum x250. 5.Megaloceraea recticornis x250. 2.Stenodema holsatum x2500 6. Trigonotylus psacsmaecolor N.S. 3.Stenodema laevigatum x250. 7. Trigonotylus ruficornis x250. 4.Notostira elongata x250. 8.Teratocoris antennatus x250. - 186 - Figure 40

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~. Teratocoris saundersi x125. 2. Teratocorls viridis x250. 3.Leptopterna dolabrata x250. 4. x250. 187 -

Plate 12 Deraeocoris ruber (Wth instar) x20 - 188 - Figure 41 a - f show micrographs of DAGOs of Vth instar nymphs, using differential interference contrast microscopy. The following structures are labelled 1 - gland sac, 2 - gland opening, 3 - secretory canal, 4 - lumen of gland, 5 - muscles attached to internal surface of gland opening. a Dicyphus globulifer (x150) - small opening, small gland sac,approximately 15 secretory canals (see fig. 42f for longitudinal section). b Blepharidopterus angulatus (x150) - medium to large opening, large gland sac, approximately 80-100 secre- tory canals (see fig. 43 c for longitudinal section). c Dicyphus pallicornis (x150) - small opening, small gland sac, approximately 20-30 secretory canals. d Monalocoris filicis (x150) - medium gland opening, large gland sac, approximately 90-100 secretory canals. e Alloeotomus gothicus (x150) - medium to large double opening, medium gland sac, approximately 40-50 secre- tory canals. f Pilgphorus perplexus (x150) - large opening, large gland sac, approximately 90-100 secretory canals (see fig. 43 a for longitudinal section) . — 1 8 9 —

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Figure 42

a shows a micrograph of a DAGO of a Vth instar nymph, using differential interference contrast microscopy. b - g show longitudinal sections through the gland sac and opening using phase contrast microscopy. The gland open- ing is always morphologically posterior to the gland sac. The following structures are labelled: 1 - gland sac, 2 - gland opening, 3 - lumen of gland, 4 - muscles attached to internal surface of gland opening, 5 - 'secretory' cells, 6 - external cuticle, 7 - cuticle lining gland sac

a Heterotoma planicornis (x150) - medium opening, medium gland sac, approximately 80-90 secretory canals. b Cyllecoris histrionicus (x250) c Asciodema obsoletum (x250) d Heterotoma planicornis (x250) - during sectioning, the dorsal layer of gland cells has become detached from the body epidermis. e Phytocoris tiliae (x250) f Dicyphus globulifer (x250) g Liocoris tripustulatus (x250) a ----0.111111.1111001■•

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Figure 43 a - f show longitudinal sections through the gland sac and opening, using phase contrast microscopy. The num- bered structures are as described in the previous figure. a Pilophorus perplexus (x250) - this specimen was near to the imaginal moult when fixed and it is possible that the absence of secretory cells is due to them having degenerated or been resorbed (there is no gland in the adult). b Leptopterna dolabrata (x250) - presence of large secretory cells may indicate high activity of the gland in this species. Under high power it is possible to . see secretory ducts from the secretory cellscpening into the lumen of gland. The cells on the ventral side of the gland appear to be larger than those on the ventral side (see also f). c Blepharidopterus angulatus (x250) - during sectioning the dorsal layer of gland cells has become detached from the body epidermis. d Dicyphus errans (x250) - dorsal layer of cells detached as in c. e (x250) - as in a, this speci- men was close to the imaginal moult. f Deraeocoris lutescens (x250) -1193 -

a b Plate 13 Deraeocoris ruber (Vth instar) x15 Note the tubular anal cone and the tufts of hairs on the lateral margins of the abdomen. - 195 - NUMERICAL TAXONOMY

Sokal and Sneath (1963) proposed two major types of classi- fication based on numerical methods - firstly, techniques of cluster analysis where taxa are arranged in a hierarchi- cal manner in the form of a phenogram (i.e. a dendrogram without phylogenetic implications), and secondly, techniques where taxa are shown as a configuration of points in p- variate Cartesian space. The advantages of numerical tech- niques are many, for example, the possibility of detailed analyses using large bodies of data, and the fact that the techniques involved can be examined stage by stage. There are, however, no optimal techniques and many workers have applied numerical methods of unknown characteristics to groups of taxa whose systematics are not fully understood. There are thus two variables and no one analysis can be said to be correct. Boratynski and Davies (1971) reviewed this problem and emphasised the importance of the compari- son and evaluation of at least some of the very many avail- able numerical techniques. In their work, they investigated the mutual relationships of 24 species of ð (Coccoidea), using mainly principal component techniques, and they related the results to the conclusions of tradit- ional taxonomic analysis. The different numerical tech- niques employed were compared and evaluated. Many plant and animal groups have been studied using numerical tech- niques, but relatively few provide comparisons of various techniques applied to the same data. Descriptions of techniques and taxa to which they have been applied can be found in Blackith and Reyment (1971) and Clifford and Stephenson (1975). More than ten years ago Sokal and Michener (1967) stated that the subject of numerical taxo- nomy still needed workers to evaluate a variety of numerical techniques on different groups of organisms.

In this study the interspecific relationships of the Vth instar nymphs of the Stenodemini, using 24 continuous, meristic characters are investigated by single linkage and average linkage clustering, and by canonical variate analy- sis. The Stenodemini was chosen for several reasons. First- ly the tribe is not too large, secondly sufficient material - 196 - had been collected and thirdly the member species appear to form a homogeneous group, having evolved as grass- feeders and with similar elongate, fusiform body shapes. The 24 characters used in the analyses were those that appear in the section on Descriptions and were measured as described on page264. The raw data for the analyses appear in statistical appendix C.

List of species investigated: Acetropis gimmerthali (Flor); Stenodema calcaratum (Fallen); S. trispinosum Reuter; S. holsatum (Fabricius); S.laevigatum (L.); Notostira elongata (Geoffroy); Megaloceraea recticornis (Geoffroy); Trigonotylus ruficornis (Geoffroy); Teratocoris antennatus (Boheman); T. saundersi Douglas and Scott; T. viridis Douglas and Scott; T.caricis Kirkaldy; Leptopterna dolabrata (L.); L. ferrugata (Fallen).

Ten specimens of each species were used in the analyses (sexes pooled) except for Teratocoris antennatus where n=6, and T. viridis, T.saundersi and T. caricis where n=4.

Clustering analyses

With the programs used, specimens with missing values for characters could be used and were denoted as -1 on the punched cards.

The initial clustering programs run dealt with only 11 species (106 OTUs), Teratocoris viridis., T. saundersi and T. caricis being added later for inclusion in the pheno- gram derived from the best clustering technique. Single linkage and average linkage programs treat each individual specimen (OTU) completely independently, no account being taken of the order of the data cards. In single linkage or nearest neighbour clustering, a new OTU is added to a cluster if it is similar to at least one of the OTUs in that cluster. In average linkage clustering, a new OTU is added to a cluster if the computed average of all

* operational taxonomic unit - 197 - similarities is above a chosen criterion level for the cluster. If the similarities are below this level, the new OTU is not admitted to the cluster.

Species OTUs A. gimmerthali 1 -10 S. calcarattun 11-20 S. trispinosum 21-30 S. holsatum 31-40 S.laevigatum 41-50 N. elongata 51-60 M. recticornis 61-70 T. ruficornis 71-80 T. antennatus 81-86 L. dolabrata 87-96 L. ferrugata 97-106

1.Single linkage clustering: program by R.G. Davies (Imperial College) Phenograms derived from correlation matrices (with data standardised and not standardised) and taxonomic distance matrices ( with data standardised and not standardised) proved unsatisfactory. Many OTUs were placed in inappro- priate clusters, but of the single linkage analyses, the phenogram derived from the taxonomic distance matrix with the data standardised gave the closest agreement with the known classification. None of the single linkage techniques were as 'good' as the average linkage clustering techniques when judged by the proportion of misclassified individuals.

2. Average linkage clustering, all matrices clustered by the weighted pair group method: program by J.C. Davis (1973) extended and amended by R.G. Davies (Imperial College) i) Phenogram derived from correlation coefficient, data not standardised. Clusters obtained: 1-10, 11-30, 31-40 including other OTUs, 41-50 excluding 46 and including other OTUs, 51-60, 61-70, 71-80, 81-86, 97-106. Number of individuals misclassified = 13 - 198 -

ii) Phenogram derived from correlation coefficient, data standardised by characters to zero means and unit standard deviations. Clusters obtained: 1-10, 11-20 excluding 12, 21-30 including 12, 31-40, 41-50, 51-60, 61-70, 71-80, 81-86, 87-96, 97-106. Number of individuals misclassified = 1

iii)Phenogram derived from distance coefficient, data not standardised. Clusters obtained: 1-10, 11-30 combined and excluding 12, 25 and 27, 31-40 excluding 32 and 38 and including 105, 41-50 excluding 41, 44 and 48, 51-60, 61-70, 71-80 exclud- ing 71, 81-86 excluding 81 and 83, 87-96 including other OTUs, 97-106 excluding 105. Number of individuals misclassified = 12

iv)Phenogram derived from distance coefficient, data standardised by characters to zero means and unit standard deviations. Clusters obtained: 1-10, 11-20 excluding 12, 21-30 includ- ing 12 and 105, 31-40, 41-50, 51-60, 61-70, 71-80, 81-86, 87-96, 97-106 ex- cluding 105. Number of individuals misclassified = 2

In all the average linkage clustering programs run, there was a good agreement with the results of conventional taxo- nomy, i.e. all the specimens of Acetropis gimmerthali were clustered together, distinct from other clusters. Mis- identifications or misplacings did occur, especially where data were not standardised. The standardisation employed produces a common spread in the data, centred about their means, where the new character form x' is computed as follows: 1 - X - X SD where x is the non-standardised variable value, x is the mean and SD is the standard deviation for that sample. - 199 -

The best fit to the known classification of the tribe appears to be given by the phenogram derived from a dis- tance coefficient matrix with the data standardised as already described, and the phenogram based on the cor- relation coefficient with data standardised. Boratynski and Davies (1971) were of the opinion that even if no serious differences are likely to arise from the use of either correlation or distance measures in clustering pro- grams, the distance form is preferable, but a decision depends partly on the kind of data being analysed and is less pronounced when continuous variables are employed. Jago (1969), using 9 continuous variables, found that clustering using a distance matrix gave a clearer separa- tion of grasshoppers of the genus Cordillacris than did clustering using a correlation matrix.

The program was rerun with an additional 12 OTUs, namely:

Species OTUs Teratocoris viridis 107-110 T. saundersi 111-114 T. caricis 115-118

The resulting phenogram is shown in Figure 44. Each OTU is not labelled by its number, but instead is shown with the number of the species to which it belongs - i.e. all . no. l's are Acetropis gimmerthali. It is immediately ap- parent that the phenogram has two major groups diverging at a taxonomic distance coefficient of 2.00*. Examination of the data reveals that those species in the left hand group (species 1,2,3,8,9,12,13,14) are small, less than 6.00mm * The scale of taxonomic distance is a relative one, and is a Euclidean distance calculated as the sum of differences over all characters between any 2 OTUs 2 k=1 (xi - xjk) d. 1j where x.u. is a measurement of the kth character on the ith row of ?ie matrix, and n = the total number of characters (Sokal, 1961). - 200 — 1N3Diii3C)) iDIMS10 Yf? evarv1 Figure 44 re e 0 O ■ - 414

O+ fo a~- J RAM G DRO R DEN STE O+ CLU E AG fo NK LI GE RA E AV

4

U

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e 2

z 01111.122 1-:

• Q b g E ~Q 1 e s . N 14 N N L Z,- ►Z ri • P~ A - 201 - in total body length, while those species in the right hand group (species 4,5,6,7,10,11) are greater than 6.00mm in total body length. In the left hand group an OTU belong- ing to group 11 (Leptopterna ferrugata) appears clustered with groups 2 and 3 (Stenodema calcaratum and S. trispinosum respectively). Examination of the specimen showed it to be malformed. S. calcaratum and S. trispinosum have not been fully separated. They are very closely related, differing in the Vth instar by having 1 and 2 metafemoral spines respectively (a unique character in the Miridae). This character is not included in the numerical analyses. The two species differ also in respect of feeding habit and the timing of the life cycle (Southwood and Leston, 1959).

Trigonotylus ruficornis is clustered with the 4 Teratocoris species and makes up the right hand side of the left major division of the phenogram. Teratocoris antennatus (group 9) has been split into 2 groups of 3 OTUs each, the left hand group being ? specimens and the right hand group d speci- mens. This subdivision into sex within a cluster occurs in two other species, Leptopterna ferrugata (group 11), where the left hand 4 are d and the right hand 5 are and in Megaloceraea-recticornis (group 7), where the left hand 5 are d and the right hand 5 are ?. The differences in size of the wing pad between sexes in some species, and other differences in body size may be responsible for the separations obtained.

Teratocoris viridis (group 12) clustered with T. saundersi and T. caricis (groups 13 and 14 respectively), the latter two appearing very similar, and this may reflect a mis- identification. The specimens of T.saundersi were collected from Juncus in the Scottish Borders (Bodesbeck Farm) and were with adults whose genitalia placed them as saundersi. However, there appeared to be some variation, and the shape of the claspers did not entirely agree with that described by. Woodroffe (1967) for this species. The dark patterns on the head and pronotum were also found to be variable. The specimens represented by groups 13 and 14 may represent one species and further study of the 6 - 202 - genitalia and other characters is required to classify this seemingly homogeneous genus.

In the major right hand group of the phenogram Notostira elongata (group 6) differs from the rest at a taxonomic distance coefficient value of 1.2. It has some morphological features which separate it from the rest of the group - the trichobothria have very weak trichomae and the trichs themselves are short. It is dangerous to ex- tract too much information from such phenograms, but it is interesting to note that on the basis of the 24 char- acters used, such techniques can agree exactly with the conventional classification.

The most suitable type of cluster analysis,on the basis of the above groups of , and using continuously variable meristic data, would appear to be one where the between- OTU matrix of distance coefficients has been calculated from data standardised by characters to zero mean and unit standard deviation, and where the matrix is clustered by the weighted pair group method. The ability of this tech- nique to further subdivide OTUs in the present study according to sex implies that the resolving power at the species level has not been exhausted.

Canonical variate analysis

Canonical variate analysis (CVA) is a multiple discriminant technique, whose main use is for examining the relation- ships between groups, the members of each group having already been established. CVA determines functions whose application to the original data, which may be transformed or standardised, maximises the observed differences between the groups. Each entity k can be considered as part of a swarm of points in multivariate cartesian space (the swarm being ellipsoidal if the variables are multi- variate normally distributed). Each entity is given a total score, calculated by summation of each of the char- acter values multiplied by an appropriate weight. The weights for different characters differ, being determined by calculation of the latent vectors of a matrix W 1B - 203 - (where W and B are sums of squares and cross product matrices) and are chosen to ensure that the variances of total scores of entities are maximised between groups as compared with the within-group variances. The underlying axes of variability produced are called the canonical variate, the 1st axis being inclined in the direction of greatest variability between the means of the k entities, the 2nd axis perpendicular to the 1st and inclined in the direction of the next greatest variability and so on for subsequent axes. CVA has been discussed by several authors (Blackith & Reyment, 1971; Clifford & Stephenson, 1975), and has been used in a great variety of plant and animal groups.

Ashton et al. (1965) studied the scapulae of a number of Anthropoidea and Prosimii, and found that the 1st 3 axes had readily interpretable biological significance. The 1st axis separated the taxa according to mode of locomo- tion (brachiators, hangers, quadrupeds etc.), the 2nd distinguished between ground and arboreal dwellers, and the 3rd axis separated man from all the rest. Eyles and Blackith (1965) investigated relationships between 4 species of the lygaeid bug Scolopostethus and their hybrids, based on 10 morphometric characters. Waloff (1966) derived meaningful results on specific and sexual variation between 2 species of mirid bug, Orthotylus virescens and O.concolor living on Sarothamnus scoparius (broom) in SE England, California and British Columbia. Examination of the clusters showed that the English specimens always had the highest value on the 1st axis, and the Canadian specimens the lowest, showing that the shape in both species had changed since their introduction to British Columbia.

In the present study 13 species of the Stenodemini were used in a CVA based on 24 morphometric characters (the same characters as used in the clustering programs) of Vth instar nymphs. The programs were written by R.G. Davies (Imperial College) and used an IBM subroutine for ex- traction of latent roots and vectors of a symmetric matrix, combined with the subroutine DIRNM of Cooley and Lohnes (1971). Unlike the clustering programs, this program - 204 - could not accept missing values, which would, strictly speaking, be inappropriate with this statistical model. Teratocoris antennatus was removed from the analysis as it had a high proportion of missing values (loss of appendages etc.). The remaining data had between 1% and 27 of values missing, these being replaced by group-character mean values. This reduced the within-group variance for these characters, but the overall effect was probably negligible.

The species studied were: Group no. Species n 1 Acetropis gimmerthali 10 2 Stenodema calcaratum 10 3 S. trispinostmn 10 4 S. holsatum 10 5 S. laevigatum 10 6 Notostira elongata 10 7 Megaloceraea recticornis 10 8 Trigonotylus ruficornis 10 9 Leptopterna dolabrata 10 10 L. ferrugata 10 11 Teratocoris viridis 4 12 T. saundersi 4 13 T. caricis 4

The table below gives the % discrimination and cumulative discrimination associated with the first 3 latent roots.

Latent root % discrimination cum % discrimination 1 45.937 45.937 2 25.821 71.758 3 10.841 82.999

The individual values for all entities on the first 3 axes have been plotted and are shown in Figures 45-47. The group mean or centroid value is represented for each group by the number denoting that group. These values are as follows: ♦ 2nd 0.1- -Tl 0 7 • 4- • ui 0.0 T ♦ • 0.2 0.3 0.4 0.5 0.7 0.8 0.9 1.0 1.1 1-2 1.3 1st

- 0.1

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-0.4- •

0I

-0•5-

-0.6- CANONICAL VAR IATES ANALYSIS (lst/2nd axes) V 3rd

0·9i 1 _ _ ~ -n G) • ~ O·SI '"

0·7 ate e~e e10Ug 0·6 N . ~ f~o~e~/g~ain o 1'~edo1l\1.nant 'j ~ f feeding '" ea 0·5 1'~edo1l\inant~'j

.t/­ Al~1J.: 0·4- -II A --

0·3 CANONICAL VARIATES ANALYSIS (lst/3rJ axes)

1st ~ 0·i ··_···· -o-~'9 1:0 1:1 1:2 1!3 .... 0·0 0·3 0·4 0·5 0·6 0·7 8 1 0

02 013 0.4 ♦ 0-6 • 0:7 0:8 0-9 1:0

Alt A 13c±A.

CANONICAL VARIATES ANALYSIS (2nd / 3rd axes) - 208 -

Species no. Group centroid values 1st axis 2nd axis 3rd axis 1 0.256 0.049 0.560 2 0.415 -0.167 0.608 3 0.366 -0.181 0.531 4 0.522 -0.213 0.704 5 0.673 -0.277 0.877 6 0.790 -0.623 0.454 7 1.193 0.033 0.490 8 0.554 0.057 0.311 9 0.618 -0.117 0.729 10 0.596 -0.067 0.668 11 0.385 -0.368 0.393 12 0.424 -0.418 0.419 13 0.405 -0.434 0.429

Statistical tests based on Wilk's lambda coefficient showed that the group means were significantly different - P<0.001.

On the plot of 1st/2nd canonical variate (fig. 45 ), Stenodema calcaratum and S. trispinosum (groups 2 and 3 respectively) are very close, as are the two Leptopterna species (groups 9 and 10). Stenodema holsatum (group 4) is fairly close to calcaratum and trispinosum, but also to the two Leptopterna spp. S. laevigatum (group 5) is quite far removed from the other three Stenodema spp. Acetropis gimmertha1i (group 1), Notostira elongata (group 6), and Megaloceraea recticornis (group 7) appear far removed from the main cluster of groups which appears to include Trigonotylus ruficornis (group 8). Examination of Figure 46 (1st and 3rd axes) shows that the latter is distinct from the central cluster of groups as well. The Teratocoris spp. cluster separately, viridis (group 11) appearing to be just distinct from saundersi and caricis (groups 12 and 13) which are very similar.

Figure 46 (1st/3rd axes) shows a basically similar picture, but with Trigonotylus, Teratocoris, Notostira and Megaloceraea distinct from the others. - 209 - Figure 47 (2nd/3rd axes) shows Trigonotylus (group 8), Teratocoris (groups 11, 12, 13), Notostira (group 6) and Stenodema laevigatum (group 5) as being distinct from the main cluster of species; and again S. calcaratum and S. trispinosum and the two Leptopterna spp. appear very close to each other.

Characters which possess absolutely high weights have more influence on the discrimination between groups than do those with absolutely low weights. It is interesting to note that total body length has a very low weight on the first 3 axes and therefore does not have a significant effect, the separation being based on shape rather than overall size. On the 1st axis, the width of the DAGO (-0.817), the length of antennal segment III (0.383) and metatarsus II (-0.215) had high weights. On the 2nd axis, the lengths of antennal segment IV (0.556), rostral segment IV (0.319), metafemur (-0.290) and head (-0.280) had high weights. On the 3rd axis the lengths of rostral segment IV (0.578), antennal segment IV (0.344), the widths of the DAGO (-0.339) vertex (-0.263) and the length of rostral segment III (0.256) had high weights. It might be possible to duplicate the results of this analysis, using fewer characters, omitting those with low weights such as total body length, metafemur width, length of rostral segments I and II etc.

In many studies using CVA it has been possible to interpret' the axes in terms of some aspect of the biology of the organisms (Ashton et al., 1965; Waloff, 1966). In the present study, the 1st axis appears to be related to the degree of elongation of the nymphs. The Stenodemini have evolved as grass-feeders (some on Carex and Juncus also) and have become elongate and fusiform as compared with other groups, this obviously having a high selective advan- tage. Some members of the tribe, however, are more elon- gate than others; for example M. recticornis is much more elongate than A. gimmerthali. A rough measure of elon- gation can be obtained by dividing the total body length by the posterior pronotal margin width (those with a low value being relatively short and squat and those with a high value being relatively elongate). The 'results are - 210 -

shown in Figure 48a. The ranges of the calculated ratio for each species are shown; the mean value is represented by the vertical bar and is projected to the horizontal axis to clarify the order. A similar ratio, this time calculated from total antennal length divided by posterior pronotal margin width, is shown in Figure 48b, using only mean values for each species. The orders obtained are basically similar to the order of species given by the 1st canonical variate, those groups with a low value being relatively short and squat, and those with a higher value being more elongate. It has not been possible to interpret the 2nd axis in terms of biological significance, but the 3rd axis may distinguish between those species that feed from flowering heads and unripe seeds of grasses and those species that feed predominantly on leaf tissue. The table below sets out the species and their feeding habits as far as is known.

Species Part of plant used Source A. gimmerthali seed V.K. Brown (personal communication) S. calcaratum buds/unripe Southwood & Leston, seed 1959 S. trispinosum flowering heads S. holsatum flowering heads/ GMcG (personal unripe seeds observation) S. laevigatum flowering heads/ Southwood & Leston, unripe seeds 1959 N. elongata leaf W.R. Dolling (pers- onal communication) M. recticornis leaf C. Gibson (1977) PhD Thesis, Oxford Tr. ruficornis leaf M.G.Morris (personal communication)' L. dolobrata leaf (early instars) C. Gibson (loc. cit.) flower (later If instars) flower/seed/leaf Southwood & Leston, (adults) 1959 L. ferrugata probably same as L. dolabrata T. viridis probably leaf T. saundersi leaf GMcG (personal observation) T. caricis leaf

see footnote on next page - 211 -

FIG. 48

A 1 11 3 8 9 134 12 10 2 6 5 7 \\\ I I — I I I

III 111f1 5 1 1 3.5 LI 4.5 5.0 •5 6.0 6.5 1 1 t 2 1 3 1 4 t 5 • 6 7 1 8 • 9 1 t 10 11 1 1 12 131

B

610

I i 3.0 [ I I 4'0 I II 1 50 I I I 3 4 10 2 11 9 8 13 12 5 6 7

A - Ratio of total body length/posterior pronotal width. The horizontal bars represent the range for each species, the mean within each range is projected on to the axis.- B - Ratio of total antennal length/posterior pronotal width - mean values for each group. 1 Acetropis gimmerthali 2 Stenodema calcaratum 3 S. trispinosum 4 S. halsatum 5 S. laevigatum 6 Notostira elongata 7 Megaloceraea recticornis 8 Trigonotylus ruficornis 9 Leptopterna dolabrata 10 L. ferrugata 11 Teratocoris viridis 12 T. saundersi 13 T. caricis - 212 -

It would be interesting to repeat the analysis using adult specimens ( although certain complications would arise in that the adults have three tarsal segments, no wing pads and no DAGO):and compare the results. This will be the subject of a future study.

* Experiments involving the cutting of grassland showed that no adverse effect to populations of Trigonotylus ruficornis resulted, and other experiments using annually mowed Dactylis and Festuca swards showed T. ruficornis as the commonest stenodemine species as well as some Notostira elongata. Species feeding on inflorescences would be unable to colonise such habitats. (M.G. Morris, personal communication). - 213 -

Plate 14 Phytocoris tiliae (IVth and Vth instars) x15 Note the distinctive mottled colouration peculiar to this genus.

- 214 - KEY TO THE SUBFAMILIES OF Vth INSTAR BRITISH MIRIDAE

1 Claws slender, with a basal projection bear- ing a flap-like or lobe-like pulvillus (figs. 55; 56 ), or claws just subapical. DAGO small and indistinct. Bothria tuber- culate Dicyphinae - Claws not as above*, never subapical. DAGO small and indistinct to large and distinct. Bothria tuberculate or not 2

2(1) Parempodia very large and flap-like (fig. 5a ). Pulvilli smal],indistinct. Dorsal sculpturing in strong polygonal pattern over entire body surface (fig. 3e,f ) . Pteridophagous Bryocorinae - Parempodia not very large ; strap-, bristle- or flap-like (figs.5c,d;7a-e;8a-e;50 ). Pulvilli small or not. Dorsal sculpturing non-polygonal. Never pteridophagous 3

3(2) Parempodia bristle-like** (figs. 5d, 53 ). Trichomae compact (figs.14d-f 15a-d) Phylinae - Parempodia strap-***, or flap-like (figs.5b,c; 7a-e; t-e) . Trichomae compact or diffuse 4

4(3) Pulvilli in live or dried specimens very dark and adpressed to claw, giving the appear- ance of a basal tooth on each claw (figs .5c; 50; 51 ). Trichomae diffuse. Parempodia strap-like***. DAGO heavily pigmented, double. Species densely pubescent Deraeocorinae - Pulvilli pale, without appearance of basal tooth on each claw. Trichomae compact. Parem- podia flap-like. DAGO heavily pigmented or not, single or double. Species densely pubescent or not 5 - 215 -

5(4) Unguitractor plate with two rows of scales. Abdominal spiracles sometimes expanded. DAGO single or with anterior bar straight, curved or weakly sinuate Orthotylinae - Unguitractor plate with three rows of scales. Abdominal spiracle openings never expanded. DAGO double or slit-like, with anterior bar often strongly sinuate Mirinae

* but see Macrotylus sp. (fig. 49) ** but see Brachyarthrum limitatum (p. 290) *** in slide mounts it may appear hair-like -216 -

Plate 15 Phytocoris varipes (Vth instar) x20 Note the distinctive mottled colouration peculiar to this genus and also the delineation (in brown) of the pharyngeal pump muscles on the head. - 217 -

Bryocorinae - key to species

1 Rostrum reaching beyond prothoracic coxae, almost to metathoracic coxae. Rostral seg- ment IV long, acute (5x as long as basal breadth) (fig. 49 ) Monalocoris filicis - Rostrum reaching to, but not beyond pro- thoracic coxae. Rostral segment IV short, squat (approx. 1.5x as long as basal breadth) (fig. 49 ) Bryocoris pteridis

Species omitted - 0

Deraeocorinae - key to species

1 Abdomen with prominent anal cone and lateral tufts of 6-14 broadly spatulate stout, black hairs. Rest of body with stout, broadly spatulate (occasionally bifurcate), black hairs. (plates 12, 13) Deraeocoris ruber - Abdomen without anal cone or lateral tuft of hairs. Body hairs simple 2

2(1) Greyish-green. Body length less than 4mm. D lutescens Colour not as above. Body length greater than 4mm 3

3(2) Body mottled reddish-brown and white. Head with longitudinal red and white markings. Metathoracic tibiae with 6 apical spines and 3 broadish red bands. Length of metatarsal segment II never more than 1.5x length of segment I. (plate 7) Alloeotomus pothicus - Colour or mottling not as above. Metathora- cic tibiae with more than 6 apical spines and without red bands. Length of metatarsal segment II always greater than 1.5x length of segment I 4

- 218 - 4(3) Antennal segment II with basal two-thirds pale, apical one-third dark and slightly incrassate. Longest body hairs 0.36-0.4amn long. Metathoracic tibiae with 12 or 13 apical spines(fig.51) Deraeocoris olivaceus - Antennal segment II uniformly dark, not in- crassate. Body hairs never longer than 0.30 mm. Metathoracic tibiae with 9 apical spines(fig.51) D scutellaris

Species omitted - Bothynotus pilosus (Boheman) (Clivenemini) See also Figure 60 for pretarsal structures.

Phylinae - key to tribes

1 Body with broad,transverse, pale band across nota, or ant mimetic. Dorsal surface with long spatulate hairs (figs .49;2d) Hallodapini - Body without pale transverse bands. Species not ant mimetic. Dorsal surface with hairs not as above. Phylini

Hallodapini - key to species

1 Pronotum parallel-sided. Abdomen often sub- globose. Metathoracic femora with cuticular sculpturing simple. Interommatidial setae spatulate Systellonotus triguttatus - Pronotum not parallel-sided Abdomen never sub-globose. Metathoracic femora with cuti- cular sculpturing specialised (fig. 2e) Interommatidial setae simple ...Hallodapus montandoni & H. rufescens - 219 - Phylini - key to species

1 Claw slender, curved and with large basal projection, from which arises a flap-like pulvillus (fig. 49 ) 2 - Claw not as above 3

2(1) Abdomen green. Body length greater than 3mm. Trichobothrial pattern 7:6 Macrotylus solitarius - Abdomen green, tinged with blue Body length not greater than 3mm. Trichobothrial pattern generally 6:5 M paykulli

3(1) Dorsal body surface with clavate hairs. (figs. 1f; 2a ) Harpocera thoracica - Dorsal body surface without clavate hairs 4

4(3) Body green with large conspicuous black spots (fig. 67 ) Oncotylus viridiflavus - Body not coloured as above 5

5(4) Parempodia flap-like, pulvilli small. Brachyarthrum limitatum - Parempodia bristle-like, pulvilli small or large 6

6(5) Dorsal surface of body with dense, broad, long, striate, apically serrate hairs (fig. 49). Body brownish-yellow. Plesiodema pinetellum - Dorsal surface of body with hairs not as above. Body colour brownish-yellow or not 7

7(6) Antennal segments I and II incrassate (fig. 52 ). DAGO with anterior bar always straight (Atractotomus) 8 - Antennal segments I and II not incrassate. DAGO with anterior bar sometimes straight 10 - 220 - 8(7) Abdomen bright red. Tibiae with distal half pale ochreous, proximal half brown. Abdomen greenish-red or yellow. Tibiae not as above 9

9(8) Body length 2.48-2.82mm. Pulvilli small, not approaching claw apices (fig. 52 ) A magnicornis Body length 2.04-2.28mm. Pulvilli larger, approaching claw apices (fig. 52 ) A mirificus

10(7) Anterior and posterior pronotal angles each with a black spot bearing a black spine (NB not all spp of Chlamydatus have such spots) Chlamydatus pullus, Pronotum not as above 11

11(10) Pulvilli large, rugulose, length greater than half the distance from the base of the pulvilli to the claw apices (fig. 52 ) 12 Pulvilli small or medium, not rugulose, length equal to half or less than half the distance from the base of the pulvilli to the claw apices 16

12(11) Pulvilli extending beyond claw apices. Wing pads extending past abdominal segment IV. Lopus, decolor Pulvilli not extending beyond claw apices. Wing pads not extending past abdominal segment IV 13

13(12) Femora and tibiae white Asciodema fieberi Femora and tibiae not white 14

14(13) Length of rostral segment IV 0.48-0.52mm. Host plant commonly Megalocoleus molliculus Length of rostral segment IV less than 0.48mm. Host plant other than above 15 - 221 -

15(14) Body uniformly green. Ratio of head. length/head width less than 0.88. Host plant Tanacetum vulgare Megalocoleus pilosus Connexivum with dark brown markings. Ratio of head length/ head width greater than 0.88. Host plants various grasses. Amblytylus nasutus

16(11) Body dark red with head and nota brown or at least with abdomen largely red 17 Body largely green,not coloured as above 24

17(16) Legs white, tibiae without strong spines or dark spots (plate 4) Orthonotus rufifrons Legs not white, tibiae with strong spines or dark spots 18

18(17) Tibiae with spines arising from large dark brown or black spots (spot up to 5x diam. of spine at base) (fig. 54 ). Host plant Psallus alnicola Tibiae with spines not arising from large spots (spot diam. small). Host plant other than above 19

19(18) DAGO single, small with anterior bar straight (fig. 25 ) Salicarus roseri DAGO double with anterior bar sinuate 20

20(19) DAGO broad (0.16-0.17mm), with dark fuscous patches laterally. Posterior margin of DAGO with strong polygonal sculpturing (fig. 54 ).(Species large, 3.84-4.21mm) Psallus betuleti DAGO not broad,or if so then without dark patches laterally or posterior polygonal sculpturing 21 21(20) Dorsal surface of abdomen with specialised hairs (fig. 54 ) P quercus Dorsal surface of abdomen without such hairs 22 - 222 22(21) Metafemoral trichobothria with strong trichomae. Bothria deeply recessed (fig.54; 15c,d) Monosynamma bohemani - Metafemoral trichobothria with weak tri- chomae. Bothria not deeply recessed 23

23(22) Body with yellow and white patches and markings. Total antennal length 1.90-2.10mm. Metatibia1 length 1.40-1.62mm. Host plant Quercus sp. Body without yellow and white patches and markings. Total antennal length 1.58-1.76mm. Metatibial length 1.20-1.36mm. Host plant Salix sp P. variabilis

24(16) Hairs of dorsal surface of head, nota and abdomen arising from small dark spots. Tibial spines never arising from conspicu- ous black spots 25 Hairs of dorsal surface of head, nota and abdomen not arising from small dark spots, or if so, then with tibial spines arising from conspicuous black spots 27

25(24) Body green with irregular red patches on lateral margins of wing pads (plate 5) . P. salicellus Body green without irregular red patches as above 26

26(25) Antennal length 1.84-2.02mm. Length of rostral segment IV . Host plant Helianthemum vulgare..Tinicephalus hortulanus Antennal length 1.68-1.77mm. Length of rostral segment IV 0.50-0.58mm. Host plant pilosella ....Hoplomachus thunbergi

27(24) Tibiae with spines arising from brown to black spots 28 Tibiae with spines not arising from brown to black spots 34 - 223 - 28(27) Tibiae with proximal ends dark brown to black, or if not, then with tibial spots very large.(fig. 54) and metafemora with one large conspicuous black spot distally on anterior margin.... (Plagiognathus sp.) 29 Tibiae with proximal ends pale. Tibial spots never very large. Metafemora with several dark spots on anterior margin, or if only one, then not large and conspicuous 31

,29(28) Femora with dark brown to black stripes on anterior and posterior margins (fig. 54 ). Plagiognathus arbustorum Femora without stripes as above 30

30(29) Dorsal body hairs black. Metatibial spines shorter than metatibial diameter. Host plants various, not usually sp. P. chrysanthemi - Dorsal body hairs pale. Metatibial spines longer than metatibial diameter. Host plants Artemisia spp. P albipennis

31(28) Metatibial spines equal to or slightly shorter than metatibial diameter....Psallus lepidus Metatibial spines longer than metatibial diameter 32

32(31) Body pale blueish-green. Body length 2.12- 2.401iuu. Host plants Althaea rosea and Verbascum spp. Campylomma verbasci Body not pale blueish-green. Body length at least 2.70mm. Hosts other than above. 33

33(32) DAGO width 0.08-0.10nuii. Antennal segment II with simple hairs. Host plants Salix spp. Psallus roseus DAGO width 0.11-0.14mm. Antennal segment II with bifurcate, occasionally trifurcate hairs. Host plants Populus spp. Sthenarus rotermundi - 224 -

34(27) Antennae with black transverse bands on proximal and distal ends of antennal seg- ment II and distal apex of antennal segment III . Phylus coryli Antennae not marked as above 35

35(34) DAGO distinctive (fig. 21 ), with anterior and lateral margins black, posterior margin pale. Tibial spines concentrated in distal half (fig. 57 ) Conostethus roseus DAGO not as shown in Figure 21 Tibial spines,if present, distributed evenly along entire length of tibiae 36

36(35) Tibiae with proximal ends fuscous. DAGO with medial constriction.(fig.23 ). Assoc- iated with Juncus. Tytthus pygmaeus Tibiae with proximal ends not fuscous. DAGO oval, without medial constriction 37

37(36) Body greyish-green. Body length 2.84-3.04mm. Metatibial length 1.38-1.44mm. Host plants Sarothamnus scoparius, Ulex europaeus. Asciodema obsoletum Body whitish-green. Body length greater than 3.2Omm. Metatibial length greater than 1.80mm. Host plant Quercus sp. Phylus melanocephalus

Species omitted - Conostethus brevis Reuter, C. griseus Douglas and Scott, Amblytylus delicatus (Perris), A. brevicollis Fieber, Tytthus geminus (Flor), Phylus palliceps Fieber, Psallus ambiguus (Fallen), P. assimilis Stichel, P. perrisi (Mulsant and Rey), P. wagneri Ossiannilsson, P. falleni Reuter, P. flavellus Stichel, P. albicinctus (Kirschbaum), P. diminutus (Kirschbaum), P. masseei Woodroffe, P. obscurellus (Fallen), P. luridus (Reuter), Plagiognathus vitellinus (Scholtz), (Fallen), C. saltitans (Fallen), C. wilkinsoni (Douglas and Scott), C. evanescens (Boheman). - 225 - Dicyphinae - key to species

1 Yellow, lateral margins of head and nota crimson. Antennae with red annuli. Claws inserted just subapically (fig. 56 ) Campyloneura virgula - Not coloured as above. Claws apical 2

2(1) Body bright red. Trichobothrial number reduced, 3:3 Dicyphus rhododendri - Body green or yellow. Trichobothrial number greater than 3:3 3

3(2) Claws curved, without strong basal projection. Pulvilli lobe-like, remote from claw (fig. 55) Macrolophus sp. - Claws nearly straight, often slightly re- curved, with a strong basal projection. Pulvilli large, flap-like approaching claw surface (fig .6 a, b ) 4

4(3) Yellowish- or greyish_-green. Head with dark anchor-shaped marking (fig. 71 ) . Dorsal body hairs sometimes arising from black spots. Trichobothrial number 4:3 5 - Pale green, greenish-red or greenish-white. Head without dark markings. Body hairs not arising from dark spots. Trichobothrial number at least 6:4 6

5(4) Body greyish-green. Body length not exceed- ing 3.Oanut. Total antennal length not exceed- ing 1.30ww. Host plant Ononis sp..Dicyphus annulatus - Body yellowish-green. Body length exceeding 3.00uuu. Total antennal length exceeding 1.30uut. Host plant Melandrium sp...... D. globulifer

- 226 - 6(4) Head and nota reddish-green. Antennal segment I entirely red. Host plant Stachys sylvatica and Lamh.im spp. Dicyphus errans - Colour not as above. Antennal segment I not entirely red 7

7(6) Metatibial length not exceeding 1.55mm. Host plant Digitalis spp. D. Pallicornis - Metatibial length greater than 1.55mm Host plant other than Digitalis 8

8(7) Metatibial length greater than 2.Oaiuu. Tricho-1 bothrial number 7:6. Host plant Epilobium hirsutism D epilobii - Metatibial length less than 2.00uiui. Tricho- bothrial number 6:5. Host plant other than above 9

9(8) Legs almost colourless D. constrictus - Legs very pale green D. stachydis

Species omitted - Macrolophus nubilus (Herrich-SchUffer), M. rubi Woodroffe. - 227 - Orthotylinae - key to tribes

1 Dorsal surface of body with short black or blackish-brown lanceolate hairs (fig. 57 ) Abdominal segment I with white transverse band. Pronotum with white transverse band along posterior margin (plates 10, 11)....Pilophorini Dorsal surface without lanceolate hairs. Abdominal segment I and posterior pronotal margin without white transverse bands 2

2(1) Rostral segments III and IV short and squat (fig.57A-D). Abdominal spiracular openings expanded, greater than diameter of tracheae (fig.57a'-D') Halticocorini Rostral segments III and IV not short and squat, or if so (Pachylops bicolor), then diameter of abdominal spiracular openings equal to diameter of tracheae (fig.57e').0rthotylini

Pilophorini - key to species

1 Dorsal surface with numerous bifurcate and trifurcate hairs Pilophorus confusus Dorsal surface with few bifurcate and tri- furcate hairs 2

2(1) Associated with Quercus and other deciduous trees (plate 10). Rostral segment IV never greater than 0.43mm. P. perplexus Associated with Pinus spp. (plate 11). Rostral segment IV equal to or greater than 0.43mm. P. cinnamopterus

Species omitted - P. clavatus (L.) - 228 - Halticocorini - key to species

1 Body very rounded and convex (fig. 69 ). DAGO absent or very small 2 - Body not rounded and convex. DAGO present, clearly visible 3

2(1) Trichobothrial number 4:3. Rostral segment IV longer than 0.20mm. Host plant Jasione montana. Strongylocoris luridus - Trichobothrial number 3:2. Rostral segment IV shorter than 0.20mm. Host plants Helianthemum vulgare, Campanula rotundifolia, and Galium verum. S. leucocephalus

3(1) DAGO with anterior bar not thickened and raised (fig. 29:1 ). Metathoracic femora not infuscate. Antennae not infuscate, slender. 4 DAGO with anterior bar thickened and raised (figs. 29:7,8 ) . Metathoracic femora largely infuscate. Antennae infuscate 5

4(3) Metathoracic tibiae not infuscate. Antennae not much longer than body. Antennal segment II less than 1.00mm long. Halticus saltator - Metathoracic tibiae with proximal two-thirds infuscate. Antennae greater than 1.1 x the body length. Antennal segment II greater than 1.00mm long. H apterus

5(3) Body length less than 3.00mm. Ratio of length of metafemur to length of metafemur trich no 3 approx. 3.5. Trichobothrial number 6:5. Pachytomella parallela - Body length greater than 3.00mm. Ratio of length of metafemur to length of metafemur trich no 3 approx. 6.5. Trichobothrial number greater than 6.5 Orthocephalus sp.

Species omitted- Halticus macrocephalus Fieber, H. luteicollis Panzer.

- 229 -

Orthotylini - key to species

1 Uniformly pale or bright green,or if not, then pale pinkish-red and occurring in salt marshes (). Trichobothrial number always 7:6, trichomae small, compact. Trichobothria never very long. Body with no distinctive features or markings. (Orthotylus sp.).15 - Not pale or bright green, or if so, at least appendages with distinctive markings. Tricho- bothrial number 7:6 or not, trichomae not always small, compact or trichobothria very long 2

2(1) Body and appendages green with proximal ends of tibiae black. Blepharidopterus angulatus - not as above 3

3(2) Antennal segments I and II greatly incrassate, with dense pubescence of black lanceolate hairs (plate 2). Heterotoma planicornis - Antennal segments I and II not greatly incrassate, or if slightly incrassate, then without black lanceolate hairs 4

4(3) Body relatively short and broad, with head, antennae and nota brown to brownish-black. Abdomen transversely banded,with lateral rows of brown or brownish-black spots.dorsally and ventrally (Heterocordylus sp.) 5 - Body more elongate. Not coloured and•marked as above 6

5(4) Trichobothrial number 6:6. Antennal segment II of ? not incrassate .(fig. 59 ). Body length 3.48-3.80mm. Host plant Sarothamnus scoparius. Heterocordylus tibialis - 230 -

Trichobothrial number 6:5. Antennal segment of ? incrassate (fig. 59 ). Body length 2.92-3.40mm. Host plant commonly Genista tinctoria, less commonly on Sarothamnus scoparius. Heterocordylus genistae

6(4) Trichomae absent, bothria flush with surface of femora 7 - Trichomae on some or all trichobothria, bothria recessed or tuberculate 8

7(6) Pale greyish-green, with three terminal abdominal segments bearing a large pair of reddish-brown patches dorsally (fig. 72 ). Dorsal body hairs mostly simple. Cyllecoris histrionicus - Green with reddish-brown and white markings. Terminal abdominal segments without patches dorsally (fig. 73 and plate 6). Dorsal body hairs black, squat, coronate (fig. 59 ) Dryophilocoris flavoquadrimaculatus

8(6) Pulvilli medium to large, extending midway to claw apices (fig. 59 ). Trichobothrial number 8:6. Shape of head distinctive (fig. 59 ) Malacocoris chlorizans , - Pulvilli small, indistinct, or if medium, trichobothrial number less than 8:6. Head not shaped as above 9

9(8) Head and nota predominantly red. Antennal segments I and II red, basal half of segment III white, apical half of III and entire IVth segment infuscate. Mecomma ambulans - Head and nota predominantly green. Antennal segments not coloured as above 10 - 231 - 10(9) Greyish-green or green. Dorsal body hairs stout, black, some bifurcate and arising from small fuscous spots. Host plant often Ulex sp. Rostral segments III and IV short and squat (fig. 57E ) Pachylops bicolor Not greyish-green or if green, then dorsal body hairs never arising from small fuscous spots. Host plant never Ulex. Rostral segments III and IV not short and squat 11

11(10) Eyes, lateral margins of pronotum and wing pads crimson. Dorsal body hairs very long (max.l. 0.22mm). Pseudoloxops coccineus Without crimson markings. Dorsal surface hairs always less than 0.22mm 12

12(11) Predominantly brownish-black. Head and pro- notum as shown in Figure 60 , pronotum expanded posteriad. Pulvilli medium or large (Globiceps) 13 Predominantly green or greenish-ochreous. Head and pronotum different from above, pro- notum not expanded posteriad. Pulvilli small or very small 14

13(12) Posterior margin of pronotum white. Terminal two abdominal segments and small patches medially on the dorsal surface of segments VI, VII and VIII fuscous... Globiceps flavomaculatus Posterior margin of pronotum concolourous with rest of pronotum. Abdominal segments with broad transverse brown or blackish- brown band. G. woodroffei

14(12) Dorsal surface of head with a diamond-shaped fuscous mark. Host plants Populus tremula, occasionally P. canescens. ... Neomecomma bilineatus Dorsal surface of head without mark as above. Associated with Carex and Juncus spp. Cyrtorhinus caricis - 232 -

15(1) Prothoracic tibiae with a group of five spines on opposite side of tibia to comb (fig. 58 ). Large species, greater than 4.00liva long. Orthotylus flavinervis - Prothoracic tibiae with a group of less than five spines. Smaller species less than 4.0011uI1 long 16

16(15) Pinkish red, occurring in salt marsh. .. 0. rubidus Green 17

17(16) Some dorsal body hairs arising from small or minute brownish-black spots 18 Dorsal body hairs never arising from brown- ish-black spots 19

18(17) Hairs arising from spots on nota only. Joint between rostral segments III and IV transverse(fig .58) 0. flavosparsus - Hairs arising from spots on entire dorsal body surface. Joint between rostral segments III and IV oblique(fig.58) 0. virescens

19(17) Setae on apical half of rostral segment IV highly curled (fig .58A, B) . 20 Setae on apical half of rostral segment IV not highly curled (fig.58C ) 21

20(19) Total antennal length 2.84-3.06mm.. 0. ochrotrichus Total antennal length 3.22-3.48mm 0. prasinus

21(19) Ratio of length of antennal segments II/III 1.45-1.61. 0. adenocarpi Ratio of length of antennal segments II/III less than 1.40. 22

22(21) Ratio of length of antennal segments II/III 1.25-1.36. Host plants Erica tetralix and Calluna spp. 0. ericetorum Ratio of length cf antennal segments II/III less than 1.25. Host plants not as above 23 - 233 - 23(22) Smaller species, body length 2.60-2.80mm. Metatibial length 1.18-1.32mm. Associated with the larger chenopods.... Orthotylus moncreaffi Larger species, body length 2.84-3.20mm. Metatibial length 1.48-1.64mm. Associated with Sarothamnus scoparius. 0. concolor

Species omitted - Fieberocapsus flaveolus (Reuter), Orthotylus fuscescens (Kirschbaum), 0. tenellus (Fallen), O. viridinervis (Kirschbaum), 0. marginalis Reuter, 0. virens (Fallen), 0. nassatus (Fabricius), 0. diaphanus (Kirschbaum), Mecomma dispar (Boheman) . - 234 - Mirinae - key to tribes

1 Strongly ant mimetic or with pronotum con- stricted anteriad and posteriad (fig. 60A & B ). Pithanini Not at all ant mimetic. Pronotum not as above 2

2(1) Entirely covered with dense, long, simple, black hairs or femora with broad medial white band. Body dark reddish-brown with white markings, shape as shown in Figure 60C. Horistini - Pubescence not as above, or if so, then body and legs strongly mottled. Femora never with broad medial white band. Body colour not as above, or if so, then shape not as in Figure60G. 3

3(2) Body elongate and fusiform. Femora never mottled (colour pattern of body often compris- ing longitudinal stripes). Trichobothrial number never 10:7 Stenodemini - Body not elongate and fusiform, or if so, then femora faintly mottled and trichobothrial number 10:7 and colour pattern of longitudi- nal stripes. Mirini

Pithanini - key to species

1 Antennal segment I black basally, paler apically. DAGO double with anterior bar sinuate. Head and pronotum as shown in Figure 60B . Not ant-like. Pithanus maerkeli - Antennal segment I unicolourous. DAGO single with anterior bar curved. Head and pronotum as shown in Figure 60C . Very ant-like. Myrmecoris gracilis

Species omitted - 0 - 235 -

Horistini - key to species

1 Femora with broad, medial white band. (Associ- ated with Melampyrtmt pratense). Capsodes flavomarginatus - Femora unicolourous or with slight pale ochreous areas 2

2(1) Dorsal body hairs very long, longest about 0.35mm. Longest tibial hairs about 0.27mm. Medial pronotal white stripe thin, width less than 0.25x the length. C. gothicus - Dorsal body hairs not very long, all distinct- ly less than 0.35mm. Tibial hairs all dis- tinctly less than 0.27mm long. Medial pro- notal stripe broad, width 0.25x length C. sulcatus

Species omitted - 0

Mirini - key to species

1 Entirely mottled, brown and white, grey and white or pale red and white, tinged with red or green. Tibiae with up to 3 distinct broad, transverse, dark bands. Antennae long and slender. Antennal segment I thickly covered with long, stout hairs (fig. 61 ). (Phytocoris sp.) 30 - Not entirely mottled, or if slightly mottled, then tibiae never with more than 2 distinct dark bands. Antennae sometimes long and slender. Antennal segment I not as above 2

2(1) Broadly oval with a large black spot on either side of midline of pro- and meso-notum. DAGO appearing as a black spot. Propleura with a single black spot adjacent to pronotal margin (fig. 74 ) (Lygus sp.) 3 Not broadly oval, or if so,then not spotted as above 4 - 236 -

3(2) Body dark green with light brown markings (fig. 74 ) . Lygus rugulipennis - Body yellowish-green without light brown markings L. maritimus

4(2) Antennae distinctly banded (fig. 75 ) Antennal segment I with reddish-brown mottling. Antennal segments II and III with broad reddish- brown bands. Antennal segment IV entirely light reddish-brown. Tibiae banded as shown in Figure 75 . Liocoris tripustulatus Antennae not as above. Tibiae not banded as above 5

5(4) Body green with small black spots on poster- ior margins of abdominal segments, distributed symmetrically about the midlines Head and nota spotted less evenly. Head with short, black longitudinal stripe medially Pantilius tunicatus - Body not spotted and marked as above 6

6(5) Antennal segment I and II slightly incrassate. Dorsal surface of body with flabellate hairs (figs. 2f; 3a ) Capsus ater - Antennal segments I and II not incrassate. Dorsal body surface without flabellate hairs 7

7(6) Head and nota with distinctive double medial red line (fig. 78 and plate 9). Trichomae absent. Megacoelum infusum - Head and nota without markings as above. Trichomae usually present 8

8(7) Trichomae very weak or absent; if very weak then body with distinctive pattern (fig. 76) 9 - Trichomae well-developed,or if weak, then body elongate with longitudinal green and white stripes down entire length 16 - 237 -

9(8) DAGO absent 10 DAGO present (but sometimes pale) 11

10(9) Body length less than 4.00mm. Host plant Juniperus sp Dichrooscytus valesianus Body length greater than 4 ..00iinti. Host plant Pinus sp. D. rufipennis

11(9) Body length never greater than 4.00mm. Body green or light brown with red markings 12 Body length always greater than 4.00mm. Body dark reddish-brown, shining, or with distinctive markings (fig. 77 ) 14

12(11) Length of rostral segment IV 0.40-0.42mm. Body light brown with distinctive red patterning (fig. 76 ). Host plant Pinus sp. Camptozygum pinastri Length of rostral segment IV 0.28-0.32mm. Body green with light red markings on wing pads or abdomen. Host plant other than Pinus sp 13

13(12) Body green with wing pad apices and femora tinged red. Host plant Viscum album. Orthops visicola Body green with faint medial red stripe down abdomen (occasionally with two lateral stripes). Host plants various Umbelliferae. 0. campestris

14(11) Body length greater than 6.5mm. Body entirely dark reddish-brown to black. Posterior margins of abdominal segments I and II yellow. Abdomen sometimes swollen. Miris striatus Body length less than 6.5mm. Not coloured as above. Abdomen never swollen 15 - 238 - 15(14) Body with distinctive patterning (fig. 77 ). Head and pronotum dull....Calocoris quadripunctatus Body not marked as above. Head and pro- notum strongly shining C. stysi

16(8) Body elongate, fusiform, with green and white longitudinal stripes..Miridius quadrivirgatus Not shaped or coloured as above 17

17(16) Hairs of dorsal surface of body stout, black, arising from small brown or black spots 18 Hairs of dorsal surface of body not short, black, or if so, then not arising from brown or black spots 20

18(17) Body dull and green. Femora and tibiae densely spotted (fig. 61 and plate 16). Adelphocoris lineolatus - Body shining green or brown Femora and tibiae not densely spotted 19

19(18) Body green. Dorsal body hairs simple. Polymerus-palustris Body dark brown, nota reddish-brown. Dorsal body hairs bifurcate, blunt.

20(17) Body small, less than 3.80mm long, squat, green and light brown. Rostral segment IV short, 0.28-0.32mm. Never with strong, black, dense pubescence...(Charagochilus sp.) 21 Body length greater than 3.80mm, squat. Rostral segment IV long, greater than 0.32mm, or if not, then very dark brown or black, or with strong,dense, black pubescence 22

21(20) Ratio of metatibial length/head width approx. 2.00. Host plant sp. Charogochilus weberi Ratio of metatibial length/head width 1.46- 1.70. Host plant Galium sp C. gyllenhali - 239 -

22(20) DAGO with strong pigmented patch posteriad and anteriad (fig. 61 ) 23 DAGO without strong pigmented patch as above 24

23(22) Body brownish-black. Polymerus nigritus Body dark green P. unifasciatus

24(22) Tibial spines arising from brown to black spots. Lygocoris contaminatus Tibial spines not arising from brown to black spots 25

25(24) Body purplish or reddish-brown dorsally 26' Body predominantly green dorsally 27

26(25) Length of metatibial spines less than half metatibial diameter. Calocoris roseomaculatus Length of metatibial spines equal to or just less than metatibial diameter. Adelphocoris ticinensis

27(25) Metatibiae with median one-third pale ochreous, basal and apical thirds brown. Calocoris alpestris Metatibiae not as above 28

28 (27) Antennal length 4.46-4.86mm. Metatibial length 2.84-3.12mm. C. norvegicus Antennal length 3.60-4.30mm. Metatibial length 2.00-2.68mm -29

29(28) Tibial spines pale. Ratio of antennal length/rostral length .2.17-2.40. Lygocoris pabulinus Tibial spines dark brownish-black. Ratio of antennal length/rostral length 1.72-1.96. Stenotus binotatus - 240 - 30(1) Head and pronotum with black stripes later- ally. Body extensively mottled greenish- brown (plate 14). Phytocoris tiliae Head and pronotum not as above. Colour as above or not 31

31(30) Associated with , especially Pinus sylvestris. P pini Not associated with conifers..other Phytocoris spp.

Species omitted - Lygus pratensis (L.), L. punctatus (Zetterstedt), L. wagneri Remane, Orthops atomarius (Meyer-DUr), 0. cervinus (Herrich-Schaffer), 0. rubricatus (Fallen), 0. kalmi (L.), Lygocoris populi Leston, L. viridis (Fallen), L. limbatus (Fallen), L. lucorum (Meyer-DUr), L. spinolai (Meyer-Diir), (Wagner), Zygimus nigriceps (Fallen), Polymerus vulneratus (Wolff), Calocoris fulvomaculatus (DeGeer), Phytocoris dimidiatus Kirschbaum, P. longipennis Flor, P. populi (L.), P. ulmi (L.), P. insignis Reuter, P. varipes Boheman.

Stenodemini - key to species

1 Metafemora with large spine or spines (fig. 19: 208,210 ) 2 Metafemora without spines 3

2(1) Metafemora with one large spine (fig.19:208) Stenodema calcaratum Metafemora with one large and one small spine (fig.19:210) S. trispinosum

3(1) Trichobothrial number 5:4. Hairs of dorsal surface of body thick, spinous, occasionally bifurcate and truncate. Acetropis gimmerthali Trichobothrial number greater than 5:4. Dorsal body hairs slender,or if not, then never bifurcate or truncate 4 - 241 - 4(3) Large species. Body length greater than or equal to 7.00mm. Antennal segment III always longer than segment II Megaloceraea recticornis Smaller species. -Body length less than 7.00nun, or if not, then antennal segment III never longer than segment II 5

5(4) Body with wide brown or green continuous or interrupted longitudinal bands. Antennal segment I not constricted subapically (figs. 61A-D; 62E-H,M,N) 6 Body without brown or green bands. Antennal segment I slightly swollen basally, constric- ted subapically (fig. 62I-L )..(Teratocoris).11

6(5) Antennae uniformly tinged red. Length of antennal segment IV less than half length of segment III. Body length usually less than 5.00mm. Trigonotylus ruficornis Antennae never uniformly tinged red. Length of antennal segment IV greater than half length of antennal segment III. Body length usually greater than 5.00mm 7

7(6) Trichobothria short (fig .19:216 , 217 ).Trichomae very weak, bothria flush with femora. Notostira elongata T richobothria not short Trichomae not weak, bothria recessed or sometimes only slightly so 8

8(7) Trichobothrial number 9:8. Body markings in form of interrupted longitudinal stripes 9 Trichobothrial number 7:6 or 7:7. Body markings in form of continuous longitudinal stripes 10

9(8) Green or greenish-yellow with dark brown markings Leptopterna dolabrata Pinkish-red with brown markings. L. ferrugata - 242 -

10(8) Ratio of head width/ metafemoral length equal to or less than 2.00. Metatibial length less than 2.50nun. Stenodema holsatum - Ratio of head width/ metafemoral length always greater than 2.00. Metatibial length greater than 2.50mm S. laevigatum

11(5) Head , nota and abdomen with longitudinal thin red stripe Teratocoris antennatus - Not as above 12

12(11) Antennal length 4.12-4.44mm. Tricho- bothrial number 7:6 T. viridis Antennal length 4.76-5.18mm. Tricho- bothrial number 9:7 or 8:7 T. saundersi & T. caricis

Species omitted - Trigonotylus psammaecolor Reuter. Plate 16 Adelphocoris lineolatus (Vth instar) x20 - 244 -

Figure 49

A Rostral segments III and IV in A - Monalocoris filicis Vth instar, bar = O.43mm B Bryocoris pteridis Vth instar, bar = O.15mm

;,l-'i'·······!·····r ...... t •••••~ Dorsal hairs in the Hallodapini ~~=.::::::;.. ..:=; ... =.;:;:::. =:;. '. ::::;: .•. =:::;;;;j~ Vth instar (x450)

B

A - Macrotylus paykulli B - M. solitarius Vth instar

L OU:::=====:::::::~ Dorsal body hairs of Plesiodema pinetellum 0--....<-'-. Vth instar (x450) - 245 -

Figure 50

Deraeocoris lutescens

D. olivaceus

D. ruber

D. scutellaris

Pretarsal structure in the Deraeocorinae Vth instar (x375) - lateral view of one half only is shown C - claw Pa - parempodium Pu - pulvillus UTP - unguitractor plate - 246 -

Figure 51

Alloeotomus gothicus

Pretarsal structures Deraeocorinae) continued

Deraeocoris olivaceus

D. scutellaris

Alloeotomus gothicus

Apical metatibial spines in some Deraeocorinae Vth instar (x175) dotted spines on other side of tibiae - 247 - Figure 52 Antennae of Atractotomus Vth instar (x80)

A. magnicornis

A. mirificus

Pretarsal structure in Atractotomus Vth instar (x375)

A. magnicornis

A. mirificus

- 248 -

Figure 53

Lopus, decolor

Megalocoleus pilosus

Megalocoleus

Amblytylus nasutus Pu Pa UTP

Asciodema fieberi

Pretarsal structures in some Phylini Vth instar (x375) C - claw Pu - pulvillus Pa - parempodium UTP - unguitractor plate

• • i -249-

Psallus alnicola - metathoracic tibia (x100)

0 • • p : • p , O

Psallus betuleti - DAGO showing poly- • •r_ . -, gonal sculpturing •4,.. 11-.),*,, posteriad 1?;,~. ~''

ti

ti - •••••••• t ` A

• A th" a . N. aa.' ā+ a 1 4 a .~+J '•:: i':• t s i J, •, ~•••:

iJ • 44

44.4444 1 ` a j N J 1 ■1 • J • 1+,J i 1 ` t ~ ~ —

Psallus quercus - Monosynamma bohemani - dorsal body hairs (x600) .metafemoral trichobothrium, deeplyzecessed (x800)

Metathoracic tibiae in Plagiognathus - Vth instar (x100)

P. albipennis

P. arbustorum

P. chrysanthemi - 250 -

Figure 55

Macrolcphus caliginosus (not a British species)

Dicyphus constrictus

D. epilobii

D. errans

D. stachydis

UTP

D. rhododendri

Pretarsal structure in the Dicyphinae Vth instar(x350) - lateral view of one half only is shown for. clarity. C - claw Pu - pulvillus Pa - parempodium UTP - unguitractor plate - 251 -

Figure 56

Dicyphus pallicornis

D. annulatus

D. globulifer

Campyloneura virgula

Pu

Pretarsal structure in the Dicyphinae - continued - 252 - Figure 57 Conostethus roseus - metathoracic tibia (x100)

Dorsal- body hairs in the 4~9 Pilophorini (x900 )

I II IV

a b' d e r I Ns

Rostral segments III and IV and abdominal spiracles in A, a' - Halticus sp. B, b' - Orthocephalus sp. C,c' - Strongylocoris leucocephalus D,d' - Pachytomella parallela E,e' - Pachylops bicolor (Rostral segments x100, spiracles X375) - 253 -

Figure 58

Orthotylus flavinervis - spines opposite comb on pro- thoracic tibiae (x175)

A B B' Rostral segments III and IV-in A Orthotylus flavosparsus B 0. virescens (lateral) B' 0. virescens (frontal) (x100)

A

B

C

Rostral segment IV in A - Orthotylus prasinus B - 0. ochrotrichus C - 0. adenocarpi Vth instar (x175)

- 254 - Figure 59 Antennae of Heterocordylus Vth instar ? (x60)

H. tibialis

H. genistae

Dorsal body hairs of Dryophilocoris flavoquadrimaculatus (x600)

A B

Malacocoris chlorizans Vth instar A - head (x60) B - metathoracic claw (x375) - 255 -

Figure 60

Head and pronoun of Globiceps woodroffei (x60)

Myrmecoris gracilis Pithanus maerkeli IVth instar Vth instar (x30) (x30)

Capsodes gothicus Vth instar (x14)

- 256 -

Antennal segment I of Phytocoris sp. (x50)

Metathoracic femur of Adelphocoris lineolatus (x30)

A

a 0

DAGOs of (A) Polymerus unifasciatus and (B) P. nigritus (x250)

A B C D Antennal segment I in the Stenodemini Vth instar (approx. x500)• A - Acetropis gimmerthali B - Stenodema calcaratum C - S. trispinosum D - S. holsatum - 257 -

Figure 62

E F G H

I L

E - S. laevigatum F - Notostira elongata G - Megaloceraea recticornis H - Trigonotylus ruficornis I - Teratocoris viridis J - T. antennatus K - T. saundersi L - T. caricis M - Teptopterna dolabrata N - L. ferrugata

Antennal segment I in the Stenodemini - continued Vth instar (approx. x500)

- 258 - Figure 63 Antennal lengths in the Stenodemini 1mm a A a a C a D

a F

a H

I

J

K

L

a M a N A Acetropis ginmerthali; B Stenodema calcaratum; C S. trispinosum; D S. holsatum; E S. laevigatun; F Notostira elongata; G Megaloceraea recticornis; H Trigonotylus ruficornis;. I Teratocoris antennatus; J T. viridis; K T. saundersi; L T. caricis; M Leptopterna dolabrata; N L. ferrugata all Vth instars - 259 -

Figure 64

4 V V J J 4 J

a

A

4 4 J V a 4 J J 4 J 4 4 J J 4 V 4 V 4 4 J J 4 4 4 J43 .4 J 4,4.4 J J J a 4444 4 , 4.4 4 p.1 44J34 44 . J J J4 ~ VV4 . 1 3 4 J;~ ~1JJ J J 4 4 4 1 44 4 : 4 4 44 44 Ni 4 3 441,4 V 4 J 1 v 4 4.31 414 4 4 4 J 1 4 4 VV 4 4 V V . 4 4J41V1J 4 4 y J V 4 4 V 4 4 4 44 4 J4 J J J .3 V 4 J 4 V 4 4 J C D

E

Diagrammatic states of a trichome around a recessed bothrium A - absent B - very weak C - weak D - marked E - strong - 2 6 0 -

Plate 17 Dicyphus pallicornis (Vth instar) x25 Note the dense covering of glaadular hairs on the host plant, foxglove (Digitalis), - 261 - DESCRIPTIONS OF BRITISH MIRID NYMPHS

The descriptions that follow are based on Vth instar nymphs unless otherwise stated in the'material examined' sections. The names and order given follow Kloet and Hincks (1964) unless otherwise stated.

A list of illustrative material follows the name, and then a description based on live, spirit-preserved and slide- mounted material. A table of diagnostic measurements for 24 body characters (where possible) is given, showing the range (x-y) and the mean (z). The range is given rather than the standard error or standard deviation because the calculations used for the latter are derived from the assumption of a normal distribution which is unlikely to be the case in the small sample numbers used, often from one locality_. The range is also more valuable to the taxonomist for identification.

Figure 65 shows the parts of the body which have been measured, and where legs and antennaecere measured, the left hand structure has been used where possible. Abbreviations used are:. w - width, 1 - length, a.w. - anterior width, p.w. - posterior width, DAGO - dorsal abdominal gland opening. The segments of the antennae, rostrum and metatarsus are numbered I to IV starting with the most proximal segment. The first antennal segment measurement includes the pedicel or basal part which fits into the antennal socket and can only be seen in slide preparations. The number of specimens measured for each structure follows the mean in cases where n is variable, otherwise it is given at the end of the measurements. (Measurements are in mm.)

The material examined is listed after the measurements, giving number of Vth instar nymphs examined, place and county names, date of collection, host plant where possible, and collector. The names of collectors are given in Appendix A, and amended county designations changed by the Local Government Act (1972) in Appendix D. All material is deposited at the British Museum of Natural History, London. - 262 -

The descriptions are divided into subfamilies as follows:

Bryocorinae pp. 264-265 Deraeocorinae pp. 266-273 Phylinae pp. 274-325 Dicyphinae pp. 326-340 Orthotylinae pp. 341-379 Mirinae pp. 380-440 - 263 -

FIG 65

A 14

13

10

11 12

18

E ~~ F

24 24 Body measurements A - whole body; B - antenna; C - rostrum; D - metathoracic leg; E - DAGO (single); F - DAGO (double). 1 total body length 2 head width 3 greatest head length in body midline 4 vertex 5 total antennal length, including pedicel 6 antennal segment I 7 TI " II 8 T1 TT III 9 TI IT IV 10 total rostral length (from bend in maxillae) 11 rostral segment I (see 10) 12 " " II 13 " 14 " " IV 15 greatest anterior pronotal width 16 " posterior " " 17 pronotal length in midline 21 tibial length 18 metathoracic wing pad length 22 tarsus I 19 metathoracic femur length 23 IT II 20 greatest femur width 24 DAGO (internal width) - 264 - Monalocoris filicis (L.) Figs. 3c,f; 19:1-3; 41d; 49

Bright green. Eyes red. Body small, squat. Rostrum with VIth segment sharp, acute, IIIrd and IVth segments indistinct, apex of rostrum reaching past prothoracic coxae, almost to mesothoracic coxae. Pronotum trapeziodal with posterior margin slightly emarginate. Dorsal surface with a few simple hairs and cuticular sculpturing as minute pimples in patterns of 4-, 5-, and 6-sided cells, often with one or two pimples in the middle of each cell. Ventral surface without sculp- turing. DAGO small, double, indistinct. Claws quite small. Pulvilli indistinct. Parempodia large, flap-like. Trichobothria 3:3, trichomae absent. Bothria flush/tuberculate.

Measurements: Total body 1. 2.20-2.48 (2.35) n=10. Head w. 0.44-0.52 (0.49) n=10, 1. 0.36-0.40 (0.38) n=10, vertex 0.22-0.30 (0.26) n=10. Antennal total 1. 1.26-1.46 (1.34) n=10, I 0.20-0.26 (0.23) n=10, II 0.40-0.52 (0.46) n=10, III 0.32-0.40 (0.35) n=9, IV 0.26-0.32 (0.30) n=9. Rostrum total 1. 0.76-0.88 (0.82) n=10. Pronotum a.w. 0.44-0.50 (0.48) n=10, p.w. 0.70-0.78 (0.74) n=10, 1. 0.36-0.40 (0.38) n=10. Wing pad 0.38-0.48 (0.43) n=10. Metafemur 1. 0.58- 0.64 (0.61) n=10, w. 0.14-0.16 (0.15) n=10. Metatibia 1. 0.80-0.86 (0.83) n=10. Metatarsus I 0.10-0.12 (0.11) n=10, II 0.22-0.30 (0.25) n=10.

Material examined: 4 ex., Herts, Harpenden, 9.VII.53, fern, TRES. 5 ex., London, Richmond, 5.VII.76, Pteridium aquilinum (L.), GMcG. 14 ex., Yorks, Malham Tarn, 27.VII.76, Pteridium aquilinum, GMcG.

Remarks: The nymphs of this species have been described by Butler (1923) and Kullenberg (1944). Bracken,P. aquilinum, and other ferns are given as host plants, the bugs feeding on the sporangia (Southwood & Leston, 1959). The unique pattern of cellular cuticular sculpturing extensively found in this and the following species appears to be unique to the subfamily, although a much reduced form of it occurs in Campyloneura virgula (Dicyphinae) and in Megalocoleus pilosus (Phylini) and Psallus sp. (Phylini)• - 265 - Bryocoris pteridis (Fallen) Figs. 5a; 19:4-5; 20:2; 49

Pale green. Base of antennal segments I and II and apical one quarter to one third of segments III and IV infuscate. Tibiae with small, pigmented band proximad. Body small, squat. Rostrum with segments III and IV small, squat, apex reaching to prothoracic coxae only. Pronotum trapeziodal. Dorsal surface hairs short and long, simple. Dorsal surface with cuticular sculpturing as minute pimples, forming cellular patterns. Ventral surface without sculpturing. DAGO small, double, with anterior bar sinuate. Claws quite small. Pulvilli indistinct. Parempodia large, flap-like. Trichobothria 4:3, trichomae absent. Bothria flush/tubercu- late.

Measurements: Total body 1. 2.44-2.80 (2.70) n=9. Head w. 0.46-0.58 (0.54) n=9, 1. 0.36-0.52 (0.41) n=9, vertex 0.28-0.32 (0.30) n=8. Antennal total 1. 1.72-1.90 (1.78) n=8, I 0.32-0.36 (0.34) n=9, II 0.42-0.60 (0.51) n=8, III 0.40-0.46 (0.43) n=8, IV 0.46-0.52 (0.50) n=8. Rostrum total 1. 0.64-0.80 (0.74) n=9. Pronotum a.w. 0.48-0.54 (0.51) n=9, p.w. 0.66-0.76 (0.71) n=9, 1. 0.34-0.42 (0.38) n=9. Wing pad 0.28-0.42 (0.34) n=9. Metafemur 1. 0.66-0.82 (0.75) n=8, w. 0.14-0.18 (0.16) n=8. Metatibia 1. 0.88-1.00 (0.95) n=9. Metatarsus 10.12-0.14 (0.14)n=9, II 0.28-0.34 (0.30) n=9.

Material examined: 10 ex., Westm, Longsleddale, 24.VII.53, fern, TRES.

Remarks: The nymphs, which have been described by Butler (1923) and Kullenberg (1944), .are very similar to the pre- ceding species, and occur on Athyrium felix-femina , Dryopteris felix-mas, other ferns and occasionally bracken, although Satchell and Southwood (1963) considered the latter unlikely as a nymphal food source in this country. They found M. filicis occurring on a wider range of ferns than B. pteridis. - 266 - Deraeocoris lutescens (Schilling) Figs. 19:6,7; 20:4; 43f; 50

Greyish-green. Eyes red. Nota mottled brown. Abdominal seg- ments V and VI with four red spots, VII, VIII, IX and X with two red spots. Dorsal surface of metathoracic femora with longitudinal red stripe. Proximal half of thoracic metatibia with two thin transverse red bands. Mesothoracic and pro- thoracic tibiae with red annulation medially. Body squat. Dorsal surface hairs very long, simple or truncate, black. Metathoracic tibiae with 13 stout spines distally. DAGO black, double, well sclerotised with anterior bar sinuate. Claw thick basally, striate. Pulvilli indistinct. Parempodia strap-or broadly hair-like. Trichobothria 7:6, trichomae very weak. Bothria flush.

Measurements: Total body 1. 3.08-3.88 (3.50) n=10. Head w. 0.70-0.84 (0.79) n=10, 1. 0.62-0.68 (0.65) n=10, vertex 0.30-0.48 (0.42) n=10. Antenna total 1. 1.60-1.78 (1.70) n=10, 10.30-0.34 (0.32) n=10, II 0.68-0.78 (0.72) n=10,III 0.28-0.34 (0.32) n=10, IV o.30-0.36 (0.33) n=10. Rostrum total 1. 1.12-1.44 (1.30) n=10. Pronotum a.w. 0.64-0.82 (0.76) n=10, p.w. 0.90-1.22 (1.12) n=10, 1. 0.48-0.58 (0.54) n=10. Wing pad o.44-0.56 (0.53) n=10. Metafemur 1.1.00- 1.12 (1.08) n=10, w. 0.20-0.26 (0.23) n=10. Metatibia 1. 1.22-1.36 (1.29) n=10. Metatarsus I 0.14-1.16 (1.15) n=10, II 0.32-0.36 (0.34) n=10.

Material examined: 8 ex., Herts, Harpenden, 7.V.54, Quercus sp., TRES. 1 ex., Berks, Silwood Park, 10.VII.76, Quercus sp., GMcG. 2 ex., Bucks, Denham, 27.VII.76, Quercus sp., GMcG. 7 ex., London, Putney Heath, 30.VII.76, Quercus sp., GMcG. 3 ex., London, Richmond Park, 10.VIII.76, Betula sp., GMcG.

Remarks: Butler (1923) described the nymphs of this species, which is predacious, and and hazel have been given as host plants (Southwood & Leston, 1959). - 267 -

Figure 66

1 2

1 12mm Bothynotus pilosus N.S. Deraeocoris lutescens (5th.)

3 4

1 51mm 1.74m

Deraeocoris olivaceus (5th.) Deraeocoris ruber (5th.) 5 6

1 06 1.30mm 4

(5th.) Deraeocoris scutellaris (5th.) A1loeotomis gothicus

- 268 -

Deraeocoris olivaceus (Fabricius) Figs. 14b; 19:8,9; 20:5; 50; 51

Pinkish-red. Antennal segment I red, segment II with base orange, apex red, incrassate, segment III with apical half red, segment IV entirely red. Head and nota brown with dark markings. Wing pads brownish-black. Abdomen mottled pink and red with lateral rows of black spots, posterior segments with black marks. Femora brown with black markings distally, posterior margins with closely spaced black spots. Tibiae with three transverse broad red bands, metatibiae with 13 stout spines distally. Tarsi black. Dorsal body hairs long, black (max.l. 0.38). Antenna with long, black hairs. DAGO strongly double, black with anterior bar sinuate. Claws thick basally, striate. Pulvilli adpressed, indistinct. Parempodia strap•or broadly hair-like. Trichobothria 7:6, trichomae present. Bothria flush.

Measurements: Total body 1. 6.66-7.58 (6.87) n=3. Head w. 1.28-1.36 (1.31) n=3, 1. 1.12-1.24 (1.18) n=3, vertex 0.70- 0.80 (0.75) n=3. Antenna total 1. 4.46 (4.46) n=1, 10.80- 0.84 (0.82) n=3, II 1.96-2.06 (2.01) n=3, III 0.94 (0.94) n=1, IV 0.64 (0.64) n=1. Rostrum total 1. 2.92-3.16 . (3.00) n=3,- I 0.80-0.90 (0.83) n=3, II 0.60-0.70 (0.63) n=3, III 0.60-0.66 (0.62) n=3, IV 0.90-0.92 (0.91) n=3. Pronotum a.w. 1.20-1.30 (1.23) n=3, p.w. 2.00-2.04 (2.01) n=3, 1. 1.02- 1.10 (1.05) n=3. Wing pad 1.00-1.06 (1.03) n=3. Metafemur 1. 2.30 (2.30) n=3, w. 0.46-0.50 (0.48) n=3. Metatibia 1. 3.00- 3.04 (3.01) n=3. Metatarsus I 0.34-0.38 (0.37) n=3, II 0.60- 0.68 (0.64) n=3.

Material examined: 1 ex., Bucks, Slough, 24.VI.55, sp, TRES. 3 ex., Herts, Bookham Common, 14.VI.56, Crataegus sp.,DL. 7 ex., Bucks, Ditton Park, 11.VI.61, Crataegus sp., GEW

Remarks: The nymphs of this partly predacious species are more hairy than those of D. scutellaris, and have longer tibial hairs. A IVth instar nymph was caught at Richmond on double-flowered hawthorn (28.V.76,GMcG). - 269 -

Deraeocoris ruber (L.) Figs. lb; 3b; 4c; 14c; 19:10,11; 20:6; 50; Pls 12; 13

Purplish-red. Antennal segment I, base of segment II black, slightly incrassate apically, segment III pale, ochreous, segment IV entirely pale red. Head and nota greyish-black with black markings. Abdominal segments I and II with hori- zontal black band, all segments with two lateral rows of black patches. Femora concolourous with body except apically, pale whitish-yellow. Tibiae with black mark proximad, longi- tudinal black stripe down entire length, otherwise pale yellow. Tarsi black. Rostrum reaching to mesothoracic coxae. Body ovate, with distinctive darkly marked tubular anal cone. Dorsal body surface with long, stout, broad, black, bi- furcate and spatulate hairs. Abdomen with lateral tufts of 6-14 black, spatulate hairs, remainder of abdominal hairs in transverse rows arising from small pigmented spots. DAGO large, double, black with anterior bar sinuate and strongly pigmented patch anteriad and posteriad. Claw thick basally, striate. Pulvilli adpressed. Parempodia strap-like, narrow. Trichobothria 7:6, trichomae present, weak. Bothria flush.

Measurements: Total body 1. 4.41-6.24 (5.84) n=10. Head w. 1.12-1022 (1.18) n=10, 1. 1.00-1.22 (1.12) n=10, vertex 0.60-0.72 (0.66) n=10. Antenna total 1. 3.26-4.40 (4.04) n=9, I 0.76-0.86 (0.80) n=10, II 1.76-2.00 (1.86) n=10, III 0.84-0.96 (0.92) n=9, IV 0.62-0.70 (0.66) n=9. Rostrum total 1. 2.32-2.44 (2.38) n=10. Pronotum a.w. 0.80-1.08 (0.97) n=10, p.w. 1.44-1.62 (1.52) n=10, 1. 0.76-0.88 (0.81) n=10. Wing pad 0.60-0.90 (0.74) n=10. Metafemur 1. 1.88-2.20 (1.97) n=10, w. 0.32-0.44 (0.39) n=10. Metatibia 1. 2.50-2.96 (2.66) n=10. Metatarsus 1 0.20-0.24 (0.23) n=10,II 0.50-0.58 (0.53) n=10.

Material examined: 2 ex., Herts, Harpenden, 15.VII.53,TRES. 3 ex., Berks, Datchet, -.VII.59, GEW. 1 ex., Notts, Radcliffe on Trent, 1.IX.76, JHM. 5 ex., Surrey, Longcross, 20.VII.77, Pinus sylvestris, GMcG. 5 ex., Bucks, Denham, 29.VII.77, sp., GMcG & MMM. - 270 -

Remarks: The nymphs of this predacious species have been described by Butler (1923), Southwood and Scudder (1956b) and Southwood and Leston (1959), and have been collected from nettles, European , bramble, broom, etc., Young instars were successfully reared on a supply of sp. aphids from a carnation. The Deraeocoris nymphs could not catch young instar nymphs of Plagiognathus arbustorum, which moved too quickly. The nymphs probed the back of the aphids between the siphunculi and inserted their rostrum . The aphids were not held by any other means and, after feeding, the rostrum was cleaned after the aphid had been pushed off the end with the prothoracic tibial combs. These combs were pushed repeatedly down the rostrum and then cleaned by a rapid flicking movement of the pro- thoracic legs. Cleaning of the antennae and the mesothoracic tibiae and tarsi by drawing through the combs then followed. The metathoracic tibiae and tarsi were then cleaned with the mesothoracic claws. Between feeding bouts, the nymphs ejected some fluids from the rostrum on to a surface, and sucked it in and out a number of times. This behaviour could serve to wash the sensory papillae at the apex of the rostrum and clean the food canal.

The eggs, nymphs, adults, life cycle and habits on hazel have been described by Viggiani (1971). Although this species is predatory, he showed that Ist instar nymphs were capable of surviving on a vegetarian diet. However, Cobben (1978) has pointed out that it was not stated whether the nymphs actually fed. - 271 -

Deraeocoris scutellaris (Fabricius) Figs. 19:12-13; 20:7; 50; 51

Antennal segments I, II, III dark brown,segment IV with red tinge. Head and nota dark brown with pale ecdysial lines. Dorsal surface of abdominal segments I, II with broad, dark brown bands, but with posterior margins pale. Posterior two abdominal segments entirely brown, rest of segments with a pair of lateral dark brown spots each in anterior half. Head squat, posterior angles of pronotum rounded. Rostrum reaching midway between pro- and meso- thoracic coxae. Dorsal surface hairs longish, simple, black (max.l. 0.26), on abdomen arising from small dark spots. Femora and tibiae with strong hairs, metathoracic tibiae with 9 stout spines distally. DAGO broad, strongly double, with anterior bar sinuate, and with pigmented patch anteriad and posteriad. Claws thick basally, pale apically. Pulvilli small, indistinct. Parempodia strap-like, narrow. Tricho- bothria 7:6, trichomae present, weak. Bothria flush.

Measurements: Total body 1. 5.49. Head w. 1.10, 1. 0.84, vertex 0.60. Antenna total 1. 3.08, I 0.50, II 1.30, III 0.66, IV 0.62. Rostrum total 1. 1.96. Pronotum a.w. 1.10, p.w. 1.46, 1. 0.74. Wing pad 0.84. Metafemur 1. 1.62, w. 0.30. Metatibia 1. 2.16. Metatarsus 10.24, II 0.60. n=1 for all measurements.

Material examined: 1 ex., Oxon, Ipsden, , GEW. - 272 -

Alloeotomus gothicus (Fallen) Figs. 4b; 5b,c; 14a; 19:14,15; 20:8; 41e; 51; Pl 7

Reddish-brown with greyish-white tinges. Antennal segments I, II brown, slightly incrassate, segments III, IV pale brown. Head with distinctive brown markings as four long- itudinal stripes tapering anteriad, ecdysial lines red, thin. Eyes red. Pronotum with anterior half brownish red, lateral margins and posterior half cream. Meso- and meta-thoracic nota red mottled with white. Abdomen mottled red with white spots and patches. Femora brownish with two indistinct pale bands apically. Tibiae white with three broad red bands and a proximal small red patch. Tarsal segments black, subequal. Body sharply oval, head acute, pronoun large with posterior angles rounded. Dorsal surface hairs long, black, simple, sometimes arising from brown or fuscous spots. Antennal segment II with stout black hairs,at right angles, segments III, IV with hairs pale, less stout. Abdomen with 6-11 long, black, stout hairs evenly spaced on each segment dorsally. Femora with strong black hairs on anterior margins only. Metathoracic tibiae with thick spines distally. DAGO strongly double, black. Claws faintly striate. Pulvilli small, indistinct. Parempodia strap-like. Trichobothria 6:5, trichomae weak, diffuse. Bothria flush.

Measurements: Total body 1. 4.32-5.49 (4.87). Head w. 0.76-1.08 (0.95), 1. 0.86-0.98 (0.90), vertex 0.50-0.56 (0.53). Antenna total 1. 2.48-2.74 (2.57), I 0.38-0.44 (0.40), II 1.20-1.32 (1.23), III 0.44-0.50 (0.46), IV 0.44- 0.50 (0.47). Rostrum total 1. 1.78-2.06 (1.93). Pro- notum a.w. 0.82-0.92 (0.88), p.w. 1.26-1.58 (1.36), 1. 0.66- 0.76 (0.71) . Wing pad 0.60-0.72 (0.65) . Metafemur 1. 1.36- 1.52 (1.41), w. 0.30-0.36 (0.'34). Metatibia 1. 1.56- 1.80 (1.64). Metatarsus 10.32-0.38 (0.35), II 0.38-0.42 (0.40). n=9for all measurements.

Material examined: 1 ex., Oxon, Swynecombe, 22.VII.64, Pinus sylvestris, GEW. 2 ex., Surrey, Witley Common, 3.VIII.65, P. sylvestris, GEW. 8 ex., Surrey, Oxshott, 17.VII.66, P.sylvestris, TRES. 1 ex., Surrey, Chobham Common, 7.VII.77, P. sylvestris, GMcG. 7 ex., Surrey, - 273 -

Longcross, 7.VII.77, P.sylvestris, GMcG. 5 ex., Berks, Silwood Park, 3.VIII.77, P. sylvestris, GMcG.

Remarks: This species, which was described in Britain in 1952 (Leston), is predacious on pine aphids of the tribe Lachinini. The DAGO is not as large and sclerotised as in Deraeocoris and the angle of the claw is less acute. - 274 -

Lopus decolor (Fallen) Figs. 4a,d; 5d; 14d,e; 19:16,17; 21:1; 53

Brownish-grey. Antennae dark, fuscous; apex of segment IV pale ochreous. Head and nota grey. Abdomen brown with green tinge. Tibiae and tarsi very dark fuscous. Rostrum reaching past prothoracic coxae, almost to mesothoracic coxae. Pronotum with angles rounded, posterior margin emarginate medially. Wing pads very long, reaching abdominal segment V. Dorsal body hairs short, simple, bifurcate or coronate (max.l . 0.028) . Ventral body hairs simple. DAGO double, pale with anterior bar sinuate. Claws thick basally, striate, tapering sharply apically. Pulvilli very large, swollen, rugulose, sometimes extending beyond claw apices. Parempodia hair-like, sometimes obscured by pulvilli. Trichobothria 6:5, trichomae present. Bothria slightly re- cessed.

Measurements: Total body 1. 2.96-3.26 (3.11). Head w. 0.68-0.74 (0.71), 1. 0.52-0.60 (0.57), vertex 0.40-0.46 (0.42). Antenna total 1. 2.18-2.50 (2.32), I 0.28-0.30 (0.29), II 0.68-0.78 (0.74), III 0.72-0.80 (0.76), IV 0.40- 0.46 (0.43). Rostrum total 1. 1.01-0.20 (1.10), I 0.27-0.34 (0.31), II 0.24-0.30 (0.27), III 0.22-0.26 (0.23), IV 0.26- 0.30 (0.28). Pronotum a.w. 0.63-0.74 (0.69), p.w. 0.80- 0.94 (0.87), 1. 0.38-0.40 (0.39). Wing pad 0.66-0.73 (0.69). Metafemur 1. 0.88-0.96 (0.93), w. 0.25-0.28 (0.26). Meta- tibia 1. 1.18-1.32 (1.25). Metatarsus I 0.10-0.12 (0.11), II 0.33-0.36 (0.35). DAGO 0.12-0.13 (0.12). n=8 in all cases.

Material examined: 1 ex., Berks, Windsor Forest, -.VII 60, GEW. 1 ex., Surrey, Chobham, -.VII.61, GEW. 11 ex., Sussex, Rye, 15.VII.77, GMcG.

Remarks: Butler (1923) briefly described the nymphs and Southwood and Leston (1959) gave common and brown bent- grasses (Agrostis sp.) as the food plant. Specimens from Rye were found among short grass mixed with Galium sp. The colour and long wing pads are distinctive features of the nymphs of this species. - 275 - Oncotylus viridiflavus (Goeze) Figs. 19:18,19; 21:2; 67

Pale olive green (sometimes bright green) with conspicuous large, black spots symmetric about body midline. Antennal segments III,IV without spots. Posterior margin of pronotum black, lateral margins of nota heavily spotted, spots merging. Tarsi, proximal tibial apices, inner wing pad margins black. Body broad, oval. Dorsal body hairs bifurcate, coronate arising from large black spots, spots sometimes merging. Ventral body surface with fewer spots, hairs simple. Pronotal hairs black (max.1. 0.17). Legs-with strong spinous hairs. DAGO large, double with anterior bar sinuate. Claws elongate, striate basally. Pulvilli large. Parempodia bristle-like. Trichobothria 8:7, trichomae strong. Bothria slightly recessed.

Measurements: Total body 1. 5.33, 5.66. Head w. 1.08, 1.10, 1. 0.88, 0.92, vertex 0.60, 0.64. Antenna total 1. 4.14, 4.30, 10.56, 0.58, II 1.54, 1.58, III 1.30, 1.42, IV 0.72, 0.74. Rostrum total 1. 1.64, 1.68, 10.38,0.40, II 0.50, 0.52, III0.34,0.36, IV 0.40, 0.42. Pronotum a.w. 0.98, 1.04, p.w. 1.24, 1.30, 1. 0.62, 0.64. Wing pad 0.80, 0.96. Metafemur 1. 1.88, 1.90, w. 0.40, 0.44. Metatibia 1. 2.80, 2.88. Metatarsus 10.22, 0.22, II 0.72, 0.76. DAGO 0.14, 0.14. n=2 in all cases.

Material examined: 3 ex., Herts, Harpenden, 26.VII.55, Centaurea nigra, TRES.

Remarks: The conspicuous large spots cE this species, previously described by Butler (1924) are distinctive. It may be that nymphs have a higher proportion of merging spots than the d nymphs. - 276 -

Figure 67

Oncotylus viridiflavus (Vth instar nymph) - 277 - Conostethus roseus (Fallen) Figs. 21:5; 57

Bright green, greenish-yellow laterally. Antennae fuscous, segment I, basal half of segment II pale ochreous. Head brownish-grey with brown markings. Eyes red. Pronotum brownish-grey with pale yellow central stripe, angles rounded. Wing pads tinged grey to reddish-grey. Abdominal segments green or greenish-yellow with two spots each, one on each lateral margin. Legs greyish-yellow. Tarsi fuscous, claws black. Dorsal body hairs short, dark, simple, be- coming stronger on legs. Ventral body hairs longish, simple. Tibiae with one or two rows of stout spines on inner face. Rostrum reaching nearly to mesothoracic coxae. DAGO small, single, constricted slightly medially, darkly pigmented laterally, with anterior bar long. Claws slender, striate basally. Pulvilli small, adpressed. Parempodia bristle- like. Trichobothria 4:4, trichs short, trichomae very weak. Bothria recessed.

Measurements: Total body 1. 2.88-3.20 (3.00). Head w. 0.66-0.74 (0.69), 1. 0.44-0.53 (0.47), vertex 0.36-0.42 (0.39). Antenna total 1. 1.36-1.56 (1.43), I 0.22- 0.24 (0.22), II 0.42-0.50 (0.45), III 0.42-0.50 (0.44), IV 0.30- 0.34 (0.31). Rostrum total 1. 1.08-1.11 (1.10), IV 0.26- 0.28 (0.27). Pronotum a.w. 0.64-0.68 (0.66), p.w. 0.81- 0.88 (0.83),1. 0.29-0.30 (0.30). Wing pad 0.50-0.52 (0.51)' n=2. Metafemur 1. 0.78-0.80 (0.79) n=3, w. 0.22-0.26 (0.23). Metatibia 1. 1.00-1.04 (1.01). Metatarsus 10.10 (0.10), II 0.34-0.36 (0.35). DAGO 0.10 (0.10). n=4 except where shown.

Material examined: 4 ex., Surrey, Stroude, 5.VI.55, TRES. 2 ex., Witley Common, -.VI.58, GEW.

Remarks: The absence of bifurcate and trifurcate dorsal hairs in unusual in the Phylini. The nymphs were previously described by Butler (1923). - 278 - Hoplomachus thunbergi (Fallen) Figs. 19:20,21; 21:6

Greenish-brown.- Antenna brown, segment IV with red tinge. Head and nota greyish-brown with posterior margins out- lined in black. Eyes red. Abdomen green with lateral rows of brown spots, terminal two segments dark brown. Posterior margins of abdominal segments suffused yellow. Tarsi fuscous, claws black. Dorsal body hairs spinous, black, bifurcate, coronate, arising from small black spots (max.l. 0.11). Ventral body surface, antennal segments II, III, IV, tibiae with simple hairs. Antennal segment I and femora with bifurcate and coronate hairs. Rostrum almost reaching meta- thoracic coxae. DAGO simple, weakly sclerotised with darkly pigmented patch anteriad and posteriad. Claws long, striate, thick basally. Pulvilli medium, lobe-like. Parerapodia bristle- like. Trichobothria 6:5, trichomae weak, compact. Bothria slightly recessed.

Measurements: Total body 1. 2.92-3.60 (3.33). Head w. 0.72-0.80 (0.76), 1. 0.62-0.70 (0.66), vertex 0.38-0.44 (0.42). Antenna total 1. 1.68-1.77 (1.72) n=5, 10.26- 0.28 (0.26), II 0.58-0.60 (0.59), III 0.46-0.50 (0.48) n=5, IV 0.36-0.41 (0.38) n=5. Rostrum total 1. 1.64-1.88 (1.76), IV 0.52-0.58 (0.55). Pronotum a.w. 0.66-0.74 (0.70), p.w. 0.90-1.00 (0.95), 1. 0.44-0.50 (0.47). Wing pad 0.55-0.66 (0.56). Metafemur 1. 0.94-1.08 (1.02), w. 0.26-0.30 (0.27). Metatibia 1. 1.36-1.54 (1.47). Metatarsus 10.12-0.14 (0.13), II 0.40-0.44 (0.42). DAGO 0.06-0.08 (0.076).

Material examined: 6 ex., Bucks, Slough, -.VI.50, GEW. 2 ex., Bucks, Longdown, -.VI.58, GEW. 3 ex., Berks, Windsor, 21.VI.61, GEW.

Remarks: Woodroffe (1953) recorded Hieracium pilosella as the food plant and Kullenberg (1944) gave a colour sketch of the Vth instar nymph. - 279 - Tinicephalus hortulanus (Meyer-Dllr) Figs. 19:22,23; 21:7

Green. Head and pronotum pale green, pronotum with broad ochreous band on posterior margin. Wing pads greenish- ochreous. Rostrum reaching past mesothoracic coxae. Dorsal body hairs biffurcate, or coronate (max.l. 0.1). Ventral $ pi body hairs/. Tibiae with dark spines, femora with stout, black hairs. DAGO large, single sclerotised with anterior bar straight and with pigmented patch anteriad and posteriad. Claws long, curved, striate basally. Pulvilli medium, rugulose. Parempodia bristle-like. Trichobothria 6:5, trichomae present, compact. Bothria recessed.

Measurements: Total body 1.3.00-3.20 (3.07). Head w. 0.74-0.80 (0.77), 1. 0.53-0.64 (0.57), vertex 0.37-0.48 (0.42). Antenna total 1. 1.84-2.02 (1.90), I 0.26-0.28 (0.27), II 0.62-0.70 (0.64), III 0.54-0.64 (0.59), IV 0.40-0.42 (0.40). Rostrum total 1. 1.56-1.68 (1.63), IV 0.43-0.50 (0.47). Pronotum a.w. 0.56-0.70 (0.66), p.w. 0.80-0.90 (0.85), 1. 0.37-0.46 (0.42). Wing pad 0.46- 0.60 (0.54). Metafemur 1. 0.99-1.06 (1.02), w. 0.29-0.36 (0.32). Metatibia 1. 1.38-1.48 (1.44). Metatarsus I 0.12- 0.13 (0.12), II 0. 39-0.42 (0.40). n=9 in all cases.

Material examined: 3 ex., Kent, Polhill Down, 9.VI.77, Helianthemum vulgare, WRD. 40 ex., Kent, Otford, 11.VI.78, H .vul g are,, WRD.

Remarks: The nymphs,which were previously described by Butler (1923),can be found in very large numbers on H. vulgare in the early part of June. - 280 -

Megalocoleus molliculus (Fallen) Figs. 19:24,25; 21:8; 53

Antennae light brown. Head and nota greenish-yellow. Wing pads, legs light brown. Abdomen greenish-yellow. Tarsi fuscous. Rostrum fuscous apically, just reaching meta- thoracic coxae. Dorsal body hairs simple and longer bifur- cate or coronate (max.l. 0.1). Antennal segments II,III,IV with simple hairs. Meso- and meta-thoracic tibiae with long coronate hairs on proximal half, stout spines on distal half. DAGO double, pale with arterior bar sinuate. Claws curved apically, thick, striate basally. Pulvilli large, rugulose, approaching claw apices. Parempodia bristle-like. Trichobothria 7:6, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 3.14-3.68 (3.49). Head w. 0.83-0.94 (0.89), 1. 0.64-0.74 (0.68), vertex 0.44-0.50 (0.46). Antennal total 1. 2.10-2.36 (2.22), 10.30-0.34 (0.32), II 0.78-0.90 (0.82), III 0.64-0.70 (0.67), IV 0.38- 0.43 (0.41). Rostrum total 1. 1.76-1.86 (1.80), IV 0.48- 0.52 (0.49). Pronotum a.w. 0.74-0.82 (0.78), p.w. 1.00- 1.06 (1.02), 1. 0.48-0.52 (0.49). Wing pad 0.55-0.70 (0.63). Metafemur 1. 1.04-1.20 (1.13), w. 0.34-0.42 (0.37). Meta- tibia 1. 1.57-1.78 (1.64). Metatarsus I 0.16-0.18 (0.16), II 0.44-0.50 (0.48). DAGO 0.13-0.14 (0.14). n=9 in all cases.

Material examined: 14 ex., Herts,Harpenden, 10.VII.53, Achillea millefolium, TRES.

Remarks: Southwood and Leston (1959) gave A. millefolium as the major host plant of this species. Kullenberg (1944) gave a colour sketch of the Vth instar nymph. - 281 - Megalocoleus pilosus (Schrank) Figs. 22:1; 53

Uniformly green. Claws and apical half of tarsus segment II black. Antennae short, segment II slightly incrassate. Wing pads reaching abdominal segment III. Rostrum reaching to mesothoracic coxae. Dorsal body hairs,including antennae and legs, strong, black. Ventral body hairs pale, simple. DAGO double, sclerotised with anterior bar sinuate. Claws curved apically, thick, striate basally. Pulvilli large, rugulose, not reaching claw tip. Parempodia bristle-like. Trichobothria 7:6?, trichomae present. Bothria recessed.

Measurements: Total body 1. 3.40-3.68 (3.58). Head w. 0.86- 0.92 (0.90), 1. 0.60-0.67 (0.64), vertex 0.40-0.50 (0.45). Antenna total 1. 2.10-2.22 (2.18), I 0.30-0.36 (0.33), II 0.74-0.80 (0.78), III 0.62-0.70 (0.65), IV 0.40-0.42 (0.41). Rostrum total 1. 1.50-1.68 (1.56), IV 0.36-0.39 (0.37). Pronotum a.w. 0.75-0.84 (0.0.79), p.w. 0.96-1.04 (1.01), 1. 0.46-0.48 (0.47). Wing pad 0.58-0.64 (0.62). Metafemur 1. 1.10-1.14 (1.13), w. 0.33-0.38 (0.36). Metatibia 1. 1.50- 1.60 (1.56). Metatarsus I 0.16 (0.16), II 0.38-0.46 (0.43). DAGO 0.14 (0.14). n=3 in all cases.

Material examined: 3 ex., Surrey, Weybridge, 21.VI.13, Tanacetum vulgare, EAB.

Remarks: Butler (1924) gave a description of the nymphs. The spines on the tibiae seem stronger and darker than those on M. molliculus and the nymph seems generally to have a thicker pubescence. - 282 - Amblytylus nasutus (Kirschbaum) Figs. 19:26,27; 22:4; 53

Green with connexiv m and apices of wing pads dark brown. Apical half of rostral segment IV, tarsus segment II and claws black. Head long, acute. Dorsal body hairs stout, bifurcate, coronate (max.l. 0.05) and shorter, simple. Ventral body surface, legs, antennal segments II, III, IV with hairs simple, occasionally bifurcate. Tibiae with stout spines. DAGO double, anterior bar highly sinuate. Claws slightly curved apically, thick, striate basally. Pulvilli large, swollen, rugulose, not reaching claw tip. Parempodia bristle-like. Trichobothria 7:6, trichomae compact, weak on mesothoracic femora. Bothria recessed.

Measurements: Total body 1. 3.36-3.90(3.59). Head w. 0.80- 0.86 (0.83), 1. 0.74-0.82 (0.77), vertex 0.40-0.46 (0.44). Antenna total 1. 2.22-2.36 (2.31) n=3, I 0.30-0.38 (0.35), II 0.78-0.84 (0.80), III 0.74-0.76 (0.75) n=3, IV 0.40-0.41 (0.40) n=3. Rostrum total 1. 1.60-1.72 (1.67), IV 0.40 (0.40). Pronotum a.w. 0.74-0.84 (0.79), p.w. 0.90-1.04 (0.98), 1. 0.42-0.44 (0.44) . Wing pad 0.50-0.58 (0.56) . Metafemur 1. 1.04-1.13 (1.10), w. 0.35-0.38 (0.36). Metatibia 1. 1.52- 1.64 (1.56). Metatarsus I 0.14 (0.14), II 0.40 (0.40). DAGO 0.10-0.12 (0.11). n=5 except where shown.

Material examined: 6 ex., Hants, Bournemouth, 30.VI.24, EAB.

Remarks: The nymphs were previously described by Butler (1924) and Southwood and Leston (1959) gave various grasses, especially Poa pratensis, as the food plants. - 283 - Macrotylus solitarius (Meyer-DUr) Figs. 22:5; 49

Green or greyish-green. Head and nota greyish-green. Wing pad apices infuscate. Abdomen bright green. Pronotum with angles rounded. Dorsal body hairs dense, long, simple, slightly tapered basally, occasionally bifurcate (max.l. 0.17). Tibiae with thick spines apically. Pronotal and abdominal hairs sometimes arising from dark spots. DAGO pale, very indistinct. Claws curved, slender with short basal projection. Pulvilli large, flap-like, remote from claw. Parempodia long, bristle-like. Trichobothria 7:6, trichomae very weak or absent. Bothria flush.

Measurements: Total body 1. 3.52-4.00 (3.79). Head w. 0.64-0.74 (0.69), 1. 0.66-0.72 (0.70), vertex 0.39-0.40 (0.40) n=4. Antenna total 1. 2.43-2.60 (2.51), I 0.30 (0.30), II 0.82-0.96 (0.91), III 0.77-0.86 (0.81), IV 0.44- 0.48 (0,46). Rostrum total 1. 1.82-2.00 (1.94), IV 0.60- 0.64 (0.62). Pronotum a.w. 0.64-0.68 (0.67), p.w. 0.82- 0.90 (0.88) n=4, 1. 0.50-0.52 (0.50). Wing pad 0.67- 0.78 (0.71). Metafemur 1. 1.32-1.48 (1.39), w. 0.29-0.34 (0.31) . Metatibia 1. 1:94-2.12 (2.05) . Metatarsus 10.16- 0.18 (0.17), II 0.48-0.56 (0.50). n=5 except where shown.

Material examined: 4 ex., Herts, Batchworth, 23.VI.60, Stachys sylvatica, DL. 1 ex., Oxon, Wychwood, 4.VIII.65, GEW. lex., Bucks, Denham, 29.VII.77, S. sylvatica, GMcG.

Remarks: The nymphs, which were described by Butler (1924), differ from Macrotylus paykulli in being larger, having a higher trichobothrial number, weak trichomae and flush bothria. However, the pretarsal structures are remarkably similar. - 284 -

Macrotylus paykulli (Fallen) Figs. 19:28,29; 22:6; 49

Bluish-green. Antennal segment I with dark markings basally. Head and nota brown with pale ecdysial lines and greenish tinges. Posterior margin of pronotum outlined in black. Abdomen green, posterior margin of segments with blue tinge. Femora and tibiae brown except for pale ochreous patches on femoral apices. Rostrum short, reaching midway between pro- and meso-thoracic coxae, with segment IV constricted apically. Dorsal body hairs simple, occasionally bifurcate on nota and posterior abdominal segment(max.l. 0.08). Pronotal hairs black, arising from minute fuscous spots. Tibiae without spines. DAGO small, oval, single, pale. Claws small, curved, withthickened basal projection. Pulvilli flap-like. Parempodia bristle-like. Trichobothria 6:5, trichomae present, weak on mesofemur. Bothria sightly recessed.

Measurements: Total body 1. 2.64-2.96 (2.75). Head w. 0.52-0.60 (0.57), 1. 0.43-0.59 (0.53), vertex 0.32-0.38 (0.36) n=8. Antenna total 1. 1063-1.76 (1.70), 10.24-0.26 (0.25), II 0.57-0.64 (0.61), III 0.50-0.56 (0.53), IV 0.30- 0.32 (0.31). Rostrum total 1. 1.03-1.23 (1.14) n=8, I 0.30-0.37 (0.33) n=7, II 0.23-0.36 (0.31) n=8, III 0.20- 0.22 (0.21) n=8, IV 0.24-0.31 (0.28). Pronotum a.w. 0.46- 0.60 (0.53), p.w. 0.68-0.76 (0.71), 1. 0.29-0.33 (0.31)0 Wing pad 0.50-0.56 (0.53). Metafemur 10 0.87-0.96 (0.93), w. 0.25-0.29 (0.27). Metatibia l0 1.24-1.44 (1.32). Metatarsus I 0.11-0.12 (0.12), II 0.33-0.38 (0.37). n=9 except where shown.

Material examined: 9 ex., Beds, Dunstable Downs, 80V.54, Ononis sp., TRES. 1 ex., Oxon, Baldhill, 17.VI.60, GEW. 1 ex., Oxon, Ipsden, 26.VI.60, GEW. 2 ex., Kent, Littlestone, 3.VI.61, GEW.

Remarks: This species, described by Butler (1923),is common- ly found on Ononis. The pretarsal structure is very similar to that of M. solitarius and the DAGO appears to be on abdominal segment III instead of the junction between III and IV, as is more usual in mirids. - 285 -

Orthonotus rufifrons (Fallen) Figs. 19:30,31; 22:7; Pis 3; 4

Dark red. Antennal segment I with red tinge and dark grey marks basally, rest of antennae pale greenish-yellow. Pronotum and wing pads with dark brown tinges. Abdominal segments with dark fuscous patches laterally. Legs pale greenish-yellow. Rostrum reaching nearly to mesothoracic coxae. Pronotum with lateral margins subparallel. Abdomen slightly swollen. Dorsal body hairs simple, occasionally bifurcate. Tibial spines weak, only apices with thickened spines. Sculpturing on wing pads and lateral abdominal areas as long randomly oriented spinules. DAGO single, pale, with anterior bar long, slightly sinuate. Claws thin, striate basally. Pulvilli medium. Parempodia bristle-like. Trichobothria 6:6, trichomae strong, compact, weaker on mesofemora. Bothria recessed. ~-

Measurements: Total body 1. 2.96-3.28 (3.12) n=10. Head w. 0.66-0.72 (0.69) n=10, 1. 0.62-0.72 (0.68) n=10, vertex 0.34-0.42 (0.37) n=10. Antenna total 1. 2.48-2.87 (2.69) n=5,I 0.30-0.36 (0.33) n=10, II 0.80-0.98 (0.90) n=10, III 0.64-0.88 (0.74) n=7, IV 0.56-0.70 (0.65) n=5. Rostrum total 1. 1.37-1.46 (1.42) n=5, IV 0.37-0.40 (0.39) n=10. Pronotum a.w. 0.60-0.64 (0.63) n=10, p.w. 0.70-0.80 (0.76) n=10, 1. 0.38-0.42 (0.41) n=10. Wing pad d 0.50-0.60 (0.54): n=6,? 0.22-0.24 (0.23) n=4. Metafemur 1. 1.08-1.20 (1.13) n=9, w. 0.22-0.26 (0.25) n=9. Metatibia 1. 1.60-1.84 (1.70) n=9. Metatarsus I 0.12-0.14 (0.13) n=7, II 0.46-0.49 (0.47) n=7. DAGO 0.08-0.09 (0.08) n=10.

Material examined: 4 ex., Berks, Little Wittenden Wood, 26.V.50, , TRES. 7 ex., Bucks, Hedgerley, -.VI.54, U. dioica, TRES. 1 (third instar) ex., Berks, Silwood Park, 3.VIII.77, Rubus sp., GMcG.

Remarks: There is marked sexual dimorphism in the nymphs of this species, the a Vth instar nymphs having long wing pads and the ? nymphs having pads half as large. This reflects the later development of the macropterous adult d and the - 286 - brachypterous adult ?. The dark red colour of the nymph becomes bright red after death. The nymphs were previously described by Butler (1923) and by Southwood and Scudder (1956b). It is possible that this species is bivoltine as a third instar nymph was caught from bramble at Silwood Park late in the season (3.VIII.77). - 287 - Harpocera thoracica (Fallen) Figs. 1e,f; 2a,b; 4e; 5e; 19:32,33; 22:8

Reddish-brown. Antennae light yellowish-brown. Head and nota reddish-brown with central white ecdysial line. Eyes red. Pronotal calli brown. Posterior margin of pronotum partly obscuring 2 or 4 small fuscous patches on anterior mesonotal margin. Abdomen reddish-brown with central pale ochreous line. Femora tinged red, especially apically. Tibiae light yellowish-brown with tarsi brown. Rostrum short, not reaching prothoracic coxae. Pronotum large. Dorsal body hairs stout, dark, clavate. Antennal segments I, II, III, femora and tibiae with clavate hairs. Antennal segment IV with bifurcate and coronate hairs. DAGO small, single with anterior bar darkly pigmented. Claws dark, thin, striate and very curved apically. Pulvilli large, rugulose. Parempodia short, bristle-like. Trichobothria 5:3, trichs short, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 4.56-5.08 (4.85). Head w. 0.96- 1.04 (1.00), 1. 0.69-0.78 (0.72), vertex 0.57-0.64 (0.61) n=6. Antenna total 1. 2.02-2.39 (2.23), I 0.34-0.44 (0.39), II 0.51-0.60 (0.56), III 0.72-0.86 (0.79), IV 0.43-,0.54 (0.47). Rostrum total 1. 1.02, 1.10 (1.06)n=2, IV 0.22-0.26 (0.24) n=8. Pronotum a.w. 0.92-1.00 (0.98), p.w. 1.46-1.64 (1.60) n=9, 1. 0.72-0.80 (0 76). Wing pad 0.72-0.83 (0.78).' Metafemur 1. 1.38-1.50 (1.46), w. 0.39-0.46 (0.42). Meta- tibia 1. 2.00-2.18 (2.08). Metatarsus 10.18-0.20 (0.19), II 0.50-0.54 (0.51). DAGO 0.05-0.06 (0.06). n=10 except where shown.

Material examined: 5 ex., Herts, Harpenden, 10.V.53, Quercus, TRES. 12 ex., Surrey, Ranmoor Common, 9.V.76, Quercus, GMcG. 3 ex., Herts, Rothamsted, 14.V.76, Quercus, GMcG. 43 ex., London, Putney Heath, 13.V. 77, Quercus, GMcG. 2 ex., Dumfs, Bodesbeck Farm, 1.VI.77, Quercus, GMcG. 2 ex., London, Wimbledon Common, 21.V.78, Quercus, GMcG,

Remarks: The nymphs were described by Butler (1923). The ' stout, black setae, truncate at apex' of Butler are infact - 288 - clavate hairs, a unique hair type in the mirid nymphs examined, and which alone enable the species to be identi- fied. The presence of these hairs and the unusual tricho- bothrial pattern and type point to the fact that this species is wrongly placed in the Phylini. The exact relationships of the species are uncertain-- however, the trichobothria show strong similarities to those in the Bryocorini. There is no sign of the specialised antennal fossae present on the apex of the second antennal segment of the adult d, these being obtained along with functional genitalia at the adult moult. - 289 -

Tytthus pygmaeus (Zetterstedt) Figs. 19:36,37; 23:2

Green. Dorsal body hairs long, dark, slightly curved, simple (max.L. 0.08 , occasionally 0.1 on posterior abdominal seg- ment). Ventral surface hairs similar. DAGO single, constric- ted medially with anterior bar slightly curved, crenulate and with small pigmented areas anteriad and laterad. Claws slender, very slightly curved. Pulvilli small, detached apically. Parempodia bristle-like. Trichobothria 7:6, trichs long, trichomae strong compact. Bothria recessed.

Measurements: Total body 1. 2.38-2.66 (2.51). Head w. 0.62- 0.66 (0.64), 1. 0.44-0.48 (0.46), vertex 0.28-0.32(0.31). Antenna total 1.2.02-2.12 (2.09), I 0.28-0.29 (0.28), II 0.58-0.63 (0.60), III 0.50-0.56 (0.53), IV 0.64-0.72 (0.68) . Rostrum total 1. 0.98-1.04 (1.01), IV 0.26-0.28 (0.27). Pronotum a.w. 0.50-0.58 (0.53), p.w. 0.60-0.70 (0.66), 1. 0.29-0.34 (0.32). Wing pad 0.37-0.40 (0.39). Metafemur 1. 0.84-0.90 (0.89) n=4, w. 0.20-0.26 (0.23). Metatibia 1. 1.14-1.24 (1.20). Metatarsus I 0.10-0.11 (0.10), II 0.35- 0.37 (0.36). DAGO 0.125 n=3. n=5 except where shown.

Material examined: 2 ex., Berks, Silwood Park, 20.VII.60, Juncus sp., TRES. 3 ex., Mddsx, Scrat✓hwood, 23.VII.60, Juncus sp., TRES.

Remarks: The immature stages and biology of this specialised mirid predator have been studied by Rothschild (1963). The nymphs feed mostly on eggs of delphacid hoppers, but also on first and second instars or dead aphids,mites and spiders. - 290 -

Brachyarthrum limitatum Fieber Figs. 19:38,39; 23:3

Grey with green tinge. Head and nota with broad, pale, medial stripe. Legs yellowish-orange. Tarsal segment II black. Dorsal body hairs long, pale, bifurcate, occasionally short, simple. Abdomen, legs and antennae with hairs becoming darker. Tibiae with thickened spines apically. Ventral cuticular sculpturing as very long, sharp spinules. DAGO single, margins black, anterior bar very dark,and with reddish patch posteriad and fuscous patch anteriad. Claws thick, striate apically. Pulvilli small. Parempodia flap-like, striate. Trichobothria 8:7, trichomae absent or very weak. Bothria flush/tuberculate.

Measurements: Total body 1. 3.20-4.16 (3.53) n=3. Head w. 0.76-0.80 (0.77) n=4, 1. 0.66 n=1, vertex 0.36-0.44 (0.41) n=4. Antenna segment I 0.34 n=1, II 0.88-0.94 (0.90) n=4, III 0.59-0.62 (0.60) n=4, IV 0.44 (0.44) n=4. Rostrum total 1. 1.36 n=1, IV 0.38 n=1. Pronotum a.w. 0.62-0.74 (0.68) n=3, p.w. 0.82-0.96 (0.88) n=4, 1. 0.46-0.48 (0.47) n=4. Wing pad 0.50-0.56 (0.54) n=4. Metafemur 1. 1.16-1.24 (1.20) n=3, w. 0.28 n=1. Metatibia 1. 1.50-1.62 (1.57) n=4. Meta- tarsus 10.12-0.13 (0.12) n=4, II 0.40-0.42 (0.41) n=4. DAGO 0.08-0.09 (0.09) n=3.

Material examined: 1 ex., Herts, Bricket Wood, 17.V.54, TRES. 2 ex., Bucks, Burnham Beeches, 4-5.VII.58, Populus,tremula, GEW. 1 ex, Surrey, Chobham, 24.VI.60, GEW.

Remarks: The flap-like parempodium is not a typical phyline character. This together with 3 rows of scales on the UTP indicates a relationship with the Mirinae. The adults have the same pretarsal structure as the nymphs, but resemble Phylus very closely and do not possess the typical mirine pronotal collar. The pretarsal structure is certainly unusual in the Phylinae. The DAGO, however, is typically phyline. The body of the nymph may appear red after death (Southwood & Le st on, 1959). - 291 -

Phylus coryli (L.) Figs. 19:34,35; 23:4

Pale green. Antennal segment II black basally and apically, III black apically. Rostral apex black. Apical half of tarsal segment II black. Rostrum reaching just past prothora- cic coxae. Dorsal body hairs bifurcate, occasionally coronate. Ventral body hairs long, thin, simple. Ventral cuticular sculpturing short, spinous. DAGO small, pale, indistinct. Claws sharply curved. Pulvilli medium. Parempodia bristle- like. Trichobothria 6:5, trichomae weak. Bothria flush.

Measurements: Total body 1. 3.76-4.04 (3.86) n=3. Head w. 0.74-0.82 (0.79)-n=4, 1. 0.66-0.68 (0.68), vertex 0.40-0.44 (0.41)-n=4. Antenna total 1. 2.42-2.76 (2.63), I 0.32-0.36 (0.35), II 0.90-1.04 (0.98), III 0.72-0.84 (0.79), IV 0.50- 0.54 (0.52) . Rostrum total 1. 1.44, 1.56 n=2, IV 0.47-0.50 (0.49). Pronotum a.w. 0.66-0.70 (0.69) n=4, p.w. 0.92-1.00 (0.94) n=4, 1..0.48-0.52 (0.49). Wing pad 0.52-0.64 (0.58). Metafemur 1. 1.16-1.28 (1.22), w. 0.26-0.33 (0.29). Meta- tibia 1. 1.78-2.04 (1.91). Metatarsus 10.12-1.14 (1.13), II 0.42-0.48 (0.46). DAGO 0.05-n=3. n=5 except where shown.

Material examined: 7 ex., Herts, Harpenden, 22.VI.54, Corylus avellana, TRES. 2 ex., Oxon, Ipsden, -.VII.61, C. avellana, GEW.

Remarks: The pattern of black marks on the aitennal segments is distinctive. Kullenberg (1944) drew a nymph of P. coryli labelled as P. melanocephalus. The nymphs of the latter species, however, lack the black markings of the antennal segments. - 292 -

Phylus melanocephalus (L.) Fig. 23:5

Pale whitish-green. Dorsal body hairs bifurcate and simple. Ventral body hairs long, thin, simple. DAGO single, weak, pale. Claws not strongly curved, striate basally. Pulvilli small to medium. Parempodia bristle-like. Trichobothria 6:6, trichomae weak. Bothria flush/tuberculate.

Measurements: Total body 1. 3.88 n=1. Head w. 0.76,0.78, 1. 0.64,0.66. Antenna total 1. 2.81,2.88, I 0.34, 0.34, II 1.11, 1.12, III 0.82,0.84, IV 0.54,0.56. Rostrum total 1. 1.50,1.60, IV 0.50. Pronotum a.wo 0.66,0.70, pow. 0.88, 1. 0.45,0.47. Wing pad 0.66. Metafemur 1. 1.30, w. 0.28. Metatibia 1. 1.92, 1.94. Metatarsus 10.13, II 0.47,0.48. n=2 except where shown.

Material examined: 2 ex., Lancs,Roudsea,Quercus,22.VI.54, Quercus sp., TRES.

Remarks; The colour of the nymphs was recorded by Southwood and Leston (1959). - 293 - Plesiodema pinetellum (Zetterstedt) Figs. 19:40,41; 23:7; 49

Brownish-yellow. Eyes red. Pronotum with posterior margins pale ochreous. Wing pads occasionally pale ochreous. Tibiae and tarsi paler than body. Dorsal body hairs slightly curved, expanded, serrate apically, (max.l. 0.05). Ventral body hairs simple. Metathoracic tibiae with thickened spines. DAGO oval, single, weakly sclerotised with anterior bar straight. Claws fairly straight, lightly striate. Pulvilli small, free apically. Parempodia bristle-like. Trichobothria 5:4, trichomae weak. Bothria flush/tuberculate.

Measurements: Total body 1. 2.34-2.65 (2.51) n=9. Head w. 0.60-0.66 (0.63) n=8, 1. 0.48-0.54 (0.52) n=9, vertex 0.28-0.34 (0.30)n=9. Antenna total 1. 1.44-1.59 (1.51) n=7, 10.20-0.22 (0.21) n=9, II 0.54-0.60 (0.57) n=9, III 0.38-0.42(0.40) n=8, IV 0.32-0.36 (0.33) n=7. Rostrum total 1. 1.20-1.23 (1.22) n=5, I 0.27-0.33 (0.29) n=4, II 0.32- 0.34 (0.33) n=4, III 0.24-0.28 (0.26) n=6, IV 0.32-0.34 (0.34) n=9. Pronotum a.w. 0.50-0.56 (0.54) n=8, p.w. 0.66- C.80 (0.76) n=76 n=8, 1. 0.33-0.37 (0.34) n=9. Wing pad 0.40-0.54 (0.48) n=9. Metafemur 1. 0.72-0.82 (0.79) n=9, w. 0.22-0.28 (0.25) n=9. Metatibia 1. 1.00-1.10 (1005)n=90 Metatarsus I 0.10-0.12 (0.11) n=9, II 0.36-0.40 (0.37) n=9. DAGO 0.04 (0.04) n=9.

Material examined: 12 ex., Bucks, Burnham Beeches, 30.VI.54, TRES. 1 ex., Surrey, Witley Common, -.VI.58, Pinus sp.,GEW.

Remarks: The nymphs of this species share an expanded, apically serrate dorsal hair structure with some Psallus spp. - 294 -

Psallus betuleti (Fallen) (S. r_pocremnus ) Fig. 54

Red. Head, nota, legs and antennae dark brown. Posterior margins of nota blackish-brown. Abdomen with two dark spots laterally per segment, segment I with broad transverse, dark bands on posterior margin, segment IV entirely dark brown. Dorsal body hairs long, bifurcate, slightly curved, becoming slightly serrate and expanded apically on abdomen. Ventral body hairs long, simple. Tibial spines striate, not longer than tibial diameter. Interommatidial setae occasionally bifurcate. DAGO large with anterior bar sinuate, pigmented laterally and with polygonal cuticular patterning on poster- ior pigmented patch. Claws short. Pulvilli small, adpressed. Parempodia bristle-like. Trichobothria 7:6, trichomae weak. Bothria flush/tuberculate.

Measurements: Total body 1. 3.84-4.21 (4.08). Head w. 0.84- 0.90 (0.87), 1. 0.60-0.70 (0.65), vertex 0.46-0.54 (0.50). Antenna total 1. 2.06-2.17 (2.12), 10.30-0.34 (0.32), II 0.86-0.92 (0.87), III 0.48-0.52 (0.50), IV 0.40-0.44 (0.41). Rostrum total 1. 1.26-1.39 (1.32), I 0.24-0.38 (0.31), II 0.30-0.36 (0.32), III 0.28-0.30 (0.29), IV 0.39-0.40 (0.40). Pronotum a.w. 0.80-0.84 (0.82), p.w. 1.08-1.14 (1.11), 1. 0.56 -0.58 (0.56). Wing pad 0.40-0.64 (0.55). Metafemur 1. .1.16 -1.21 (1.18), w. 0.34-0.39 (0.37). Metatibia 1. 1.62- 1.74 (1.66). Metatarsus I 0.16 -0.18 (0.17), II 0.42-0.48 (0.45). DAGO 0.16-0.17 (0.17). n=4 in all cases.

Material examined: 5 ex., Herts, Harpenden, 14.VI.54, Betula sp., TRES.

Remarks: Kullenberg (194 4) gave a colour sketch of a Vth instar nymph of this species. Two of the above five speci- mens were parasitised by the larvae of a braconid , which filled almost the entire abdominal space. The poly- gonal sculpturing posterior to the DAGO is very similar to that seen in the Bryocorinae. - 295 - Psallus variabilis (Fallen) (S. Hylopsallus)

Reddish-brown. Antennal segments pale brown. Head,nota, femora brown. Abdomen red with 2 faint lateral spots per segment. Tibiae pale brown. Dorsal body hairs bifurcate, serrate, often parallel-sided, sometimes tapering apically or slightly expaided apically. Ventral body hairs simple. Tibiae with stout, short spines arising from pale spots. DAGO double, anterior bar strongly sinuate and with very faint polygonal sculpturing posteriad. Claws striate. Pulvilli small. Parempodia bristle-like. Trichobothria 7:6, trichomae weak. Bothria flush/tuberculate.

Measurements: Total body 1. 2.92-3.19 (3.08). Head w. 0.70 -0.76 (0.72), 1. 0.52-0.58 (0.55), vertex 0.34-0.44 (0.41). Antenna total 1. 1.58-1.76 (1.67) n=6, I 0.22-0.24 (0.23), II 0.60-0.68 (0.64), III 0.42-0.48 (0.45), IV 0.30-0.38 (0.34) n=6. Rostrum total 1. 1.12-1.22 (1.16), I 0.21-0.28 (0.24), II 0.30-0.32 (0.31), III 0.23-0.28 (0.26), IV 0.34- 0.36 (0.35). Pronotum a.w. 0.62-0.70 (0.66), p.w. 0.86- 0.96 (0.89), 1. 0.42-0.48 (0.45). Wing pad 0.49-0.54 (0.52). Metafemur 1. 0.88-0.96 (0.93), w. 0.28-0.34 (0.31). Meta- tibia 1. 1.20-1.36 (1.29). Metatarsus I 0.12-0.14 (0.13) n=6, II 0.38-0.42 (0.41) n=6. DAGO 0.11 (0.11). n=7 except where shown.

Material examined: 7 ex., Herts, Harpenden, 28.V.53, Quercus sp.; TRES.

Remarks: This species is recorded from oak, although it is believed to be 'confined to sallow and aspen' (Southwood & Leston, 1959). Butler (1923)• described the nymphs and recorded them from oak, and Kullenberg(1944) gave a colour sketch of a Vth instar nymph. - 296 - Psallus quercus (Kirschbaum) (S. Asthenarius) Fig. 54

Dark red. Antennae, head, nota, legs brownish-red, nota with posterior margins very dark brownish-red. Terminal abdominal segment dark, segment I with dark transverse band, all segments with two faint spots laterally. Coxal margins fuscous. Dorsal body hairs broad, serrate, striate, expanded apically, or occasionally short, simple. Ventral body hairs simple. Tibial hairs bifurcate,. coronate and short, simple. DAGO single, broad, with anterior bar not very sinuate, and with faint polygonal sculpturing posteriad and dark pigmented bands laterally. Claws thick, striate. Pulvilli small, Parempodia bristle-like. Trichobothria 7:6, trichomae weak. Bothria flush/tuberculate.

Measurements: Total body 1. 2.76-3.60 (3.20). Head w. 0.70- 0.77 (0.74), 1. 0.51-0.66 (0.59), vertex 0.36-0.43 (0.40). Antenna total 1. 1.76-2.28 (2.02), 10.24-0.28 (0.26), II 0.68-0.88 (0.77), III 0.48-0.66 (0.58), IV 0.36-0.46 (0.42). Rostrum total 1. 1.30-1.53 (1.42), I 0.28-0.39 (0.33) n=5, II 0.33-0.38 (0.36) n=5, III 0.27-0.36 (0.33) n=5, IV 0.34- 0.44(0.40). Pronotum a.w. 0.65-0.71 (0.68), p.w. 0.88- 0.97 (0.92), 1. 0.42-0.48 (0.45). Wing pad 0.50-0.60 (0.55). Metafemur 1. 0.92-1.08 (0.99), w. 0.26-0.34 (0.31). Meta- tibia 1. 1.30-1.58 (1.42). Metatarsus 10.12 (0.12) n=5, II 0.39-0.43 (0.41) n=5. DAGO 0.10-0.12 (0.11) . n=6 except where shown.

Material examined: 7 ex., Lancs, Devil's Gallop, 113rL54, Quercus sp., TRES.

Remarks: The dorsal surface hairs of this not very common species are very similar to those of Plesiodema pinetellum. No live specimens of this species were available for examin- ation, but it is probable that the colouring of the nymphs is largely red. - 297 - Psallus alnicola Douglas and Scott (S. Psallus s.s.) Figs. 19:42,43; 24:1; 54

Reddish-green. Antennal segment I with two pigmented spots, rest of antennae pale. Head and nota reddish-green. Abdomen bright red. Legs pale with large fuscous spots on tibiae and less so on femora. Antennal segments I, II twice as thick as segments III, IV. Rostrum reaching mid-way between meso- and meta-thoracic coxae. Dorsal body hairs bifurcate, trifurcate, coronate, striate, arising from dark spots on head, nota, legs. Ventral body hairs simple. Mesonotal wing pad sculpturing as minute dots or pimples. DAGO pale,single with anterior bar.pale, slightly sinuate. Claws striate basally. Pulvilli small to medium. Parempodia bristle-like. Trichobothria 6:5, trichomae compact. Bothria slightly recessed.

Measurements: Total body 1. 2.52-3.04 (2.77). Head w. 0.68- 0.74 (0.71), 1. 0.48-0.58 (0.52), vertex 0.36-0.40 (0.37). Antennal total 1. 1.64-1.78 (1.70) n=8, I 0.22-0.24 (0.23), II 0.64-0.77 (0.71), III 0.42-0.50 (0.45), IV 0.30-0.32 (0.31) n=8. Rostrum tctall. 1.20-1.28 (1.24) n=8, I 0.32 n=1, II 0.32 n=1, III 0.24 n=2, IV 0.32-0.35 (0.34) n=8. Pronottnn a.w. 0.60-0.66 (0.62), p.w. 0.80-0.87 (0.81), 1. 0.37-0.44 (0.41). Wing pad 0.48-0.56 (0.51). Metafemur 1. 0.86-0.92 (0.88), w. 0.28-0.35 (0.30). Metatibia 1. 1.24- 1.44 (1.32). Metatarsus 10.10-0.12 (0.11), II 0.34-0.39 (0.36). DAGO 0.06-0.07 (0.06). n=9 except where shown.

Material examined: 9 ex., Mddsx, Hampton Court, 11.VII.76, Alnus glutinosa, GMcG.

Remarks: This species, described by Butler (1923) and Southwood and Leston (195 9) is confined to alder. - 298 - Psallus lepidus (Fieber) (S. Psallus s.s.)

Green. Antennal segments I, II greenish-yellow with a few pale spots, segments III, IV without spots. Head, nota yellow. Abdomen with a few pale spots. Tibiae with many blackish-brown spots. Dorsal body hairs long, pale, bifur- cate, trifurcate and occasionally expanded apically. Ventral hairs long, simple. Antennal segment I with two stout spines arising from black spots. Tibiae with strong spines, longer than tibial diameter, arising from black spots. Mesothoracic wing pad with cuticular sculpturing as minute raised dots or pimples. DAGO pale, single with anterior bar almost straight. Claws not stout or slender. Pulvilli medium. Parempodia bristle-like. Trichobothria 7:6, trichomae present. Bothria slightly recessed.

Measurements: Total body 1. 3.24-3.68 (3.42). Head w. 0.72- 0.86 (0.78), 1. 0.50-0.60 (0.55), vertex 0.39-0.46 (0.42). Antennal total 1. 1.80-2.14 (1.98), I 0.24-0.28 (0.26), II 0.70-0.86 (0.78), III 0.46-0.56 (0.52), IV 0.40-0.44(0.41). Rostrum total 1. 1.00-1.18 (1.09), I 0.34 n=1, II 0.30 n=1, III 0.24 n=1, IV 0.28-0.32 (0.29). Pronotum a.w. 0.63-0.76 (0.69), p.w. 0.92-1.04 (0.96), 1. 0.42-0.50 (0.56). Wing pad 0.50-0.60 (0.56). Metafemur 1. 0.95-1.08 (1.02), w. 0.30-0.34 (0.32). Metatibia 1. 1.32-1.52 (1.43). Meta- tarsus 10.12-0.13 (0.12), II 0.40-0.42 (0.41). DAGO 0.08- 0.09 (0.08). n=4 except where shown.

Material examined: 4 ex., Lancs, Old Park, 22.VI.54, Fraxinus excelsior, TRES.

Remarks: The nymphs of this species were previously described by Butler (1923). - 299 -

Psallus roseus (Fabricius) (S. Psallus s.s.)

Greyish-green. Femora with a few pigmented spots. Rostrum reaching mid-way between meso- and meta-thoracic coxae. Dorsal body hairs bifurcate, trifurcate (max.l. 0.14). Ventral body hairs simple. Antennal segment I with two stout spines. Interommatidial setae long, simple. Tibiae with stout spines arising from fuscous spots. DAGO pale,single with anterior bar weakly sinuate. Claws thick basally, striate. Pulvilli medium. Parempodia bristle-like. Tri- chobothria 8:7, trichomae often associated with a pigmented spot. Bothria recessed.

Measurements: Total body 1. 2.84-3.28 (2.99). Head w. 0.70- 0.72 (0.71),1. 0.49-0.52 (0.51), vertex 0.35-0.39 (0.38) n=3. Antenna total 1. 1.64-1.86 (1.78) n=3, I 0.24(0.24), II 0.70-0.76 (0.72), III 0.32-0.52 (0.44) n=3, IV 0.38-0.40 (0.39) n=3. Rostrum total 1. 1.16-1.26 (1.21) n=3, IV 0.34- 0.36 (0.35) n=3. Pronotum a.w. 0.60-0.63 (0.62) n=3, p.w. 0.78-0.83 (0.80) n=3, 1. 0.36-0.40 (0.39). Wing pad 0.42- 0.54 (0.50) n=3. Metafemur 1. 0.90-0.98 (0.94), w. 0.26- 0.30 (0.28). Metatibia 1. 1.32-1.44 (1.39). Metatarsus I 0.12 (0.12), II 0.36-0.39 (0.37). DAGO 0.08-0.09 (0.09). n=4 except where shown.

Material examined: 4 ex., Westm, Helsingford , 31.VI.54, Salix sp., TRES.

Remarks: One of the four specimens above was parasitised by the larva of a braconid wasp. Butler (1923) described the nymphs and Southwood and Leston (1959) recorded greyish- green as being the general colouration of the nymphs. - 300 -

Psallus varians (Herrich-Sch1ffer) (S. Psallus s.s.)

Antennal segment I with red tinge, segment IV entirely red, rest of antennae light fuscous yellow. Head, nota often with yellow and white patches and markings. Eyes white. Abdomen red with terminal segment fuscous. Femora, tibiae with dark spots. Rostrum reaching past prothoracic, nearly to mesothoracic coxae. Dorsal body hairs pale, bifurcate, trifurcate, occasionally coronate, arising from dark spots on head, nota, abdominal segments I and II. Ventral body hairs simple. Tibial spines black arising from dark spots. DAGO pale, single with anterior bar pale, sinuate. Claws striate basally. Pulvilli medium. Parempodia bristle-like. Trichobothria 8:7, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 3.20-3.68 (3.44). Head w. 0.76- 0.80 (0.77), 1. 0.56-0.62 (0.59), vertex 0.36-0.43 (0.39). Antenna total 1. 1.90-2.10 (2.03), I 0.24-0.28 (0.27), II 0.76-0.84 (0.80), III 0.52-0.60 (0.58), IV 0.37-0.40 (0.39). Rostrum total 1. 1.28-1.38 (1.34), IV 0.37-0.41 (0.39). Pronotum a.w. 0.67-0.72 (0.69), p.w. 0.90-1.02 (0.98), 1. 0.46-0.52 (0.49). Wing pad 0.50-0.56 (0.54). Metafemur 1. 1.06-1.14 (1.11), w. 0.33-0.36 (0.35). Metatibia 1. 1.40-1.61 (1.56). Metatarsus I 0.13-0.14 (0.14), II 0.42- 0.46 (0.45). DAGO 0.10-0.11 (0.11). n=5 in all cases.

Material examined: 7 ex., Lancs, High Newton, 1.VI.54, Quercus sp., TRES. 2 ex., Berks, Windsor, 12.V.61, GEW. 1 ex., Bucks, Slough, 21.V.61, GEW.

Remarks: The nymphs of this species,which are abundant on oak, have been previously described by Butler (1923) and Southwood and Leston (1959). - 301 - Psallus salicellus (Herrich-SchIffer) P1 5

Bright green. Antennal segment I with red spot ventrally, segment IV with red tinge, rest of antennae green. Head with red tinges around antennal sockets and on lateral margins. Wing pad lateral margins with irregular red mark- ings and apices dark green. Abdomen with pale yellow spots. Apical half of tarsal segment II, claw, rostral segment IV black. Rostrum reaching to mesothoracic coxae. Dorsal body hairs bifurcate, trifurcate, occasionally coronate, expand- ing slightly apically. Ventral body hairs simple. Nota with dark hairs arising from yellowish- brown spots. Tibial spines long, stoutish, arising from black spots. DAGO pale red, single with anterior bar slightly sinuate. Claws slightly striate. Pulvilli medium. Parempodia bristle-like. Trichobothria 7:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 1.54. Head w. 0.74, 1. 0.56, vertex 0.40. Antenna total 1. 1.88, I 0.24, II 0.76, III 0.48, IV 0.40. Rostrum total 1. 1.32, IV 0.38. Pronotum a.w. 0.68, p.w. 0.86, 1. 0.40. Wing pad 0.54. Metafemur 1. 0.96, w. 0.24. Metatibia 1. 1.38. Metatarsus I 0.12, II 0.40. n=1 in all cases.

Material examined 1 ex., Berks, Silwood Park, 3.VIII.77, Rubus sp., GMcG.

Remarks: Southwood and Leston (1959) recorded this species from bramble, hazel, sallow, apple and alder and described it as being partly predacious on mites, such as Bryobia praetiosa and other small animals. - 302 -

Atractotomus magnicornis (Fallen) Figs. 19:44, 45; 24:2; 52

Dark reddish-green. Antennal segments I, II red, segments III, IV pale ochreous. Head, nota brownish-green. Eyes red. Wing pad apices brown to dark brown. Abdomen dark reddish- green, becoming transversely banded after death, segment I with transverse brown band, connexivum with fuscous patches ventrally. Femora pale with a few defined black spots. Antennal segments I, II incrassate. Dorsal body hairs stout, long, black, bifurcate, arising from dark red to black spots and thinner, shorter, bifurcate, occasionally trifurcate, not arising from pigmented spots. Ventral body hairs long, black, simple. Antennal segment I with two stout, blunt hairs. Antennal and interommatidia1 setae simple. Meta- thoracic tibiae with stout, long, black spines arising from black spots, and thinner long, black hairs arising from extremely faint spots. Cuticular sculpturing as minute raised dots. Posterior margins of head and pronotum with dorsal sculpturing in clumps broken up by a network of non- sculptured cuticle. DAGO oval, single with anterior bar straight, lying in centre of black patch. Claws straight, striate basally. Pulvilli small. Parempodia bristle-like. Trichobothria 7:6, trichomae compact. Bothria recessed.

Measurements: Total body 1. 2.48-2.82 (2.64). Head w. 0.64, -0.72 (0.68), 1. 0.48-0.53 (0.51), vertex 0.30-0.36 (0.33), Antenna total 1. 1.57-1.72 (1.65) n=7, 10.22 -0.24 (0.22), II 0.58-0.66 (0.62), III 0.40-0.48 (0.44)n=7, IV 0.33-0.38 (O.36)n=7. Rostrum total 1. 1.02-1.18 (1.09), III 0.24-0.28 (0.26)n=7, IV 0.30-0.32 (0.31). Pronotum a.w. 0.53-0.60 (0.56),p.w. 0.70-0.80 (0.75),1. 0.34-0.40 (0.39). Wing pad 0.42-0.56 (0.48). Metafemur 1. 0.72-0.92 (0.86), w. 0.20-0.28 (0.24). Metatibia 1. 1.06-1.22 (1.16). Meta- tarsus I 0.10-0.12 (0.11), II 0.36-0.37 (0.36). DAGO 0.06 (0.06). (Antennal segment II w. 0.08-0.10 (0.09)).n=8 except where shown.

Material examined: 12 ex., Bucks, Slough, -.VI.53, Picea sp., GEW. 1 ex., Herts, Harpenden, 21.VI.53, Picea sp., TRES. 8 ex., Berks, Silwood Park, 3.VII.77, Picea sp., GMcG. - 303 - Atractotomus mali (Meyer-DUr) Figs. 19:46,47; 24:3, 52

Red to reddish-brown. Antennal segments I, II red, III, IV pale with yellow tinge. Head, nota, femora brownish-red. Eyes red. Nota with pale medial ecdysial lines. Abdomen bright red, segment I with brown transverse band, terminal two segments brownish-red. Coxal margins black. Tibiae with distal halves brownish-red, proximal halves pale. Tarsi pale. Rostrum reaching to mesothoracic coxae. Dorsal body hairs pale, trifurcate, parallel-sided, slightly expanded apically (max.1. 0.13). Ventral hairs simple. Antennal seg- ments I, II incrassate, with dark to black, slightly lanceolate hairs, segments III, IV with hairs pale. Meta- tibial spines stout, long. DAGO oval, single with anterior bar straight. Claws smooth. Pulvilli medium. Parempodia bristle-like. Trichobothria 7:6, trichs long, trichomae compact. Bothria recessed.

•Measurements: Total body 1. 2.60-3.00 (2.78). Head w. 0.70- 0.76 (0.73), 1. 0.52-0.58 (0.57), vertex 0.36-0.40 (0.38). Antenna total 1. 1.53-1.58 (1.56)n=7, 10.22-0.24 (0.24), II 0.56-0.60 (0.59), III 0.38-0.40 (0.39)n=7, IV 0.33-0.36 (0.34)n=7. Rostrum total 1. 1.08-1.20 (1.12), III 0.24- 0.26 (0.25)n=5, IV 0.31-0.36 (0.33). Pronotum a.w. 0.59- 0.70 (0.65), p.w. 0.79-0.86 (0.81), 1. 0.39-0.44 (0.41). Wing pad 0.46-0.54 (0.50). Metafemur 1. 0.80-0.94 (0.89), w. 0.30-0.36 (0.32). Metatibia 1. 1.12-1.26 (1.19). Meta- tarsus 10.11-0.12 (0.12), II 0.36-0.36 (0.37). DAGO 0.04 (0.04). (Antennal segment II w. 0.12-0.16 (0.14)). n=8 except where shown.

Material examined: 8 ex., Bucks, Slough, 3.VII.54, Crataegus sp., TRES. 4 ex., Herts, Harpenden, 12.VI.54, Fraxinus excelsior, TRES. 2 ex., London, Hampstead Heath, 25.VI.76, Crataegus sp., GMcG.

Remarks: Butler (1923) described the nymphs of this partially predacious species. As well as the above mentioned trees, it is also found on apple (Southwood & Leston, 1959). - 304 - Atractotomus sp. (mirificus Woodroffe,1971) Figs. 13; 19:48,49; 52

Pale green or greenish-yellow. Antennal segments I, II red, segments III, IV pale yellow. Head, nota brownish-green. Eyes red. Nota with posterior margins dark. Abdomen greenish- yellow with faint transverse orange marks. Tibiae, tarsi, rostral segment IV fuscous. Antennal segment I, II incrass- ate, segments III, IV subequal. Dorsal body hairs shortish, black, bifurcate or simple, long on pronotum and posterior abdominal segments. Ventral body hairs black, simple. Anten- nal segments I, II with hairs thort, black, simple, occasion- ally bifurcate, segments III, IV with hairs pale, short, simple. Anterior angles of pronotum with one strong black hair directed anteriad. Abdominal hairs in transverse rows. Metafemora with two stout black hairs on anterior margins. Tibiae with stout black spines. DAGO small, single with anterior bar straight. Claws stout, striate basally. Pulvilli large, rugulose, extending nearly to claw apices. Parem- podia bristle-like. Trichobothria 8:7, trichs long, trichomae present. Bothriaiecessed.

Measurements: Total body 1. 2.04-2.20 (2.17)n=6. Head w. 0.60 -0.62 (0.61) 1. 0.42-0.46 (0.45), vertex 0.28-0.30 (0.29). Antenna total 1. 1.23-1.32 (1.29), I 0.20 (0.20), II 0.30- 0.50 (0.48), III 0.30-0.32 (0.30), IV 0.29-0.32 (0.31). Rostrum total 1. 0.92-1.00 (0.95), IV 0.26-0.29 (0.28). Pro- notum a.w. 0.48-0.57 (0.53) n=6, p.w. 0.64-0.71 (0.68) n=6, 1. 0.30-0.33 (0.31). Wing pad 0.39-0.47 (0.43). Meta- femur 1. 0.65-0.70 (0.68), w. 0.20-0.22 (0.21). Metatarsus 10.10 (0.10), II 0.30-0.31 (0.30). (Antennal segment II w. 0.08 (0.08)). n=7 except where shown.

Material examined: 7 ex., Surrey, Longcross, 20.VII.77, Pinus sylvestris, GMcG.

Remarks: These specimens possibly represent the nymphs of A. mirificus Woodroffe, 1971. However, adult 66 found with them show some puzzling features and it is possible that this is an undescribed species. The dissection of one 6 showed that the aedeagus was partly like A. mirificus, being - 305 - strongly 'S'-shaped, but half like A. magnicornis in having the secondary gonopore at the apex. Another d, however, showed the typical mirificus shape (Woodroffe, 1971). It is possible that A. mirificus is a variarqe species and more collecting is required from various local- ities from Pinus and Picea in order to establish the degree of variation in the shape of the aedeagus.in the two species. The antennae of the adults collected were more like those of A. magnicornis than A. mirificus. The nymphs appear distinct from those of A. unicornis and A. mali. - 306 - Plagiognathus albipennis (Fallen) (S. Poliopterus) Figs. 19:50,51; 24:4 ; 54

Green. Antennal segment I, base of segment II fuscous. Rostral segment IV fuscous. Apices of wing pads fuscous. Tarsi fuscous. Rostrum extending to mesothoracic coxae. Dorsal body hairs long, pale, bifurcate, trifurcate, becoming darker and stouter on posterior abdominal segments (max.1.0.14). Antennal segment I with three spines. Pro- thoracic femora with one stout black spine arising from a black spot on anterior distal surface. Meso- and meta- thoracic femora with two stout spines arising from well- defined black spots distally. Meso- and meta-thoracic tibiae with two rows of very strong spines, longer than tibial diameter, arising from large black spots and one shorter, weaker row with fewer spines arising from small, ill-defined black spots. DAGO double with anterior bar sinuate. Claws long, straight. Pulvilli small. Parempodia bristle-like. Trichobothria 7:6, trichomae compact. Bothria recessed.

Measurements: Total body 1. 2.56-2.80 (2.64)._ Head w. 0.64- 0.68 (0.66), 1. 0.47-0.52 (0.49), vertex 0.32-0.36 (0.34). Antenna total 1. 1.76-1.89 (1.83), 10.22-0.24 (0.24), II 0.58-0.64 (0.61), III 0.52-0.58 (0.55), IV 0.42-0.44 (0.43). Rostrum total 1. 1.00-1.16 (1.11), IV 0.30-0.32 (0.31). Pronotum a.w. 0.57-0.60 (0.54), p.w. 0.74-0.80 (0.77), 1. 0.33-0.36 (0.35). Wing pad 0.42-0.50 (0.47). Metafemur 1. 0.84-0.90 (0.86), w. 0.28-0.31 (0.29) . Metatibia 1. 1.16- 1.24 (1.20). Metatarsus 10.11-0.12 (0.12), II 0.35-0.38 (0.36)n=9. DAGO 0.07-0.08 (0.07). n=10 except where shown.

Material examined: 12 ex., Essex, Flatford, 12.V.53, Artemisia sp., TRES. 8 ex., :Dorset, Portland, 20.VIII.77, A. absinthium, RH.

Remarks: The nymphs of this bivoltine species have been described by Butler (1923), Puchkov and Puchkova (1956) and Southwood and Leston (1959). The latter authors mention that the nymphs can be distinguished from the other species in the genus by their lack of black hairs. Some black hairs occur, but fewer than in the other species of the genus. - 307 - Southwood and Blackith (1960) recognised only one of the new species designated by Wagner as contained in P. albipennis, namely P. collinus. They discussed the considerable intraspecific variation in P. albipennis and the effect of host plants. The use of multivariate morphometrics suggested that the differences between the forms of P. albipennis on different plants could be attributed to the selection of certain genotypes on each host, together with a modifying effect from the host plant's environment. It is not certain whether the species examined were P.albipennis or P. collinus, but were probably the former. - 308 -

Plagiognathus arbustorum (Fabricius) (S. Plagiognathus s.s.) Figs. 19:52,53; 24:5; 43c; 54; 68

Yellowish-green to green. Antennal segment I, basal half of segment II, basal quarter of segment I1I brown, zest of antenna pale. Head, nota light brownish-green. Apices of wing pads dark brown in late Vth instar. Anterior margins of metafemora with dark brown to black longitudinal stripes extending to distal apices, posterior margins with one •large and one small brown or black spot and faint longitudi- nal brown stripe. Meso- and pro-thoracic femora similarly marked. Tibiae with proximal apices black. Rostrum reaching to metathoracic coxae. Dorsal body hairs sparse, short, black, bifurcate or trifurcate. Ventral body hairs slender, simple. Tibiae with two rows of stout spines, just longer than tibial diameter, arising from black spots. DAGO pale, single with anterior bar straight. Claws slender,curved. Pulvilli medium. Parempodia bristle-like. Trichobothria 7:6, trichomae compact. Bothria slightly recessed.

Measurements: Total body 1. 3.52-4.22 (3.70). Head w. 0.72- 0.78 (0.76), 1. 0.60-0.66 (0.63), vertex 0.36-0.40 (0.37). Antenna total 1. 2.20-2.36 (2.26), I 0.30-0.32 (0.32), II 0.78-0.90 (0.81), III 0.64-0.72 (0.70), IV 0.44-0.46 (0.45). Rostrum total 1. 1.28-1.44 (1.36), I 0.24-0.34 (0.29), II 0.36-0.44 (0.39), III 0.24-0.32 (0.28), IV 0.38-0.42 (0.40). Pronotum a.w. 0.66 -0.76 (0.71), p.w. 0.80-0.90 (0.86), 1. 0.42-0.48 (0.44) . Wing pad 0.40-0.68 (0.53) . Metaf emur 1. 1.14-1.26 (1.20), w. 0.28-0.36 (0.31). Metatibia 1. 1.56- 1.78 (1.66). Metatarsus I 0.12-0.14 (0.13), II 0.42-0.50 (0.47). n=10 in all cases.

Material examined: 7 ex., Bucks, Hedgerley,-.VI.54, TRES. 11 ex., Herts, Harpenden, 1.VII.55, TRES. 12 ex., London, Hampstead Heath, 25.VI.77, Urtica dioica, GMcG. 2 ex., Surrey, Chobham, 7.VII.77, GMcG. 3 ex., Perths, Kinloch Rannoch, 26.VI.78, U. dioica, GMcG.

Remarks: The nymphs of this very abundant, partially preda- cious species have been described by Butler (1923), - 309 -

Figure 68

Plagiognathus arbustorum (Vth instar nymph) - 310 -

Van Dinther (1953) and Southwood and Scudder (1956b)and are readily identifiable by the longitudinal black stripes on the metathoracic femora. Specimens were reared from IIIrd instar to maturity on a diet of Myzus certes (Walker) from carnation . - 311 -

Plagiognathus chrysanthemi (Wolff) (S. Plagiognathus s.s.) Figs. 19:54,55; 24:6; 54

Green. Antennal segment I, base of segment II with dark markings, rest of antenna pale or light brown. Head , nota green. Rostral segment IV fuscous. Tibiae with dark brown- black spots, and with proximal apices fuscous. Tarsi fuscous. Dorsal body hairs black. Tibial spines shorter than tibial diameter, arising from fuscous spots. DAGO double with anterior bar sinuate. Claws striate basally, not very slender. Pulvilli small to medium. Parempodia bristle-like. Trichobothria 7:5, trichomae compact. Bothria recessed.

Measurements: Total body 1. 2.84-3.41 (3.15). Head w. 0.66- 0.78 (0.73), 1. 0.52-0.60 (0.58), vertex 0.32-0.44 (0.41). Antenna total 1. 1.98-2.21 (2.10), 10.26-0.30 (0.27), II 0.66-0.77(0.73), III 0.58-0.66 (0.63), IV 0.41-0.47(0.45). Rostrum total 1. 1.28-1.52 (1.39), 10.30-0.52 (0.38), II 0.30-0.39 (0.36), III 0.26-0.30 (0.28), IV 0.34-0.40 (0.37). Pronotum a.w. 0.52-0.74 (0.68), p.w. 0.74-0.88 (0.84), 1. 0.34-0.42 (0.40). Wing pad 0.39-0.62 (0.52). Metafemur 1. 1.00-.109 (1.04), w.0.28-0.32 (0.30). Metatibia 1. 1.36- 1.53 (1.46). Metatarsus 10.11-0.12 (0.12), II 0.40-0.44 (0.42). n=10 in all cases.

Material examined: 4ex., Bucks, Slough, -.VI.53, GEW. 14ex., Herts, Great Field, 25.VI.53 and 2.VII.53, TRES. 1 ex., Berks, Silwood Park, 3.VII.77, GMcG. 14 ex., Perths, Kinloch Rannoch, 26.VI.78, GMcG.

Remarks: The black dorsal hairs separate this species from P. albipennis,and the lack of femoral stripes from P. arbustorum. At Kinloch Rannoch many Vth instar nymphs were taken running on upland heath, dominated by Erica tetralix, Thymus drucei and Galium saxatile. The species is recorded as being especially common in dry areas of waste ground on Senecio sp., Achillea millefolium and Medicago lupulina (Southwood & Leston ,1959), and was described by Butler (1923) and Southwood and Scudder (1956b). - 312 - Chlamydatus pullus (Reuter) (S. Euattus) Figs. 19:58,59; 25:1; 69

Green. Antennal segment I with basal half fuscous, apical half pale, segment II with basal half fuscous becoming pale apically, segments III, IV pale or very lightly in- fuscate. Eyes red bordered with white. Apical half of tarsal segment II and claws entirely black. Rostrum reaching to mesothoracic coxae, apex of segment IV black. Metathoracic femora slightly expanded. Dorsal body hairs black, long, stout, bifurcate, or less stout, simple (max. 1. 0.11). Ventral body hairs slender, pale , simple. Pro- notal anterior and posterior angles with a black spot bearing a stout black hair (the anterior spots being the larger). Anterior margins of femora with black hairs arising from small black spots (one on prothoracic femur, two on meso- and meta-thoracic femur). Tibiae with two rows of stout black hairs arising from black spots. Metathora- cic tibial hairs as long as tibial diameter. DAGO double, pale with anterior bar sinuate. Claws striate, very slightly curved. Pulvilli medium. Parempodia bristle-like. Tricho- bothria 7:6, trichs long, trichomae appearing as small dark spots. Bothria well-recessed.

Measurements: Total body 1. 2.12-2.28 (2.19). Head w. 0.65- 0.68 (0.66), 1. 0.40-0.44 (0.42), vertex 0.33-0.38 (0.35). Antenna total 1. 1.22,1.23 n=2, I 0.20 (0.20), II 0.36- 0.38 (0.37), III 0.32, 0.32, IV 0.33,0.34. Rostrum total 1. 0.85-0.90 (0.86), IV 0.22-0.23 (0.22). Pronotum a.w. 0.54 (0.54), p.w. 0.66-0.70 (0.68), 1. 0.29-0.32 (0.31). Wing pad 0.28-0.30 (0.29). Metafemur 1. 0.67-0.68 (0.68), w. 0.27-0.28 (0.28). Metatibia 1. 0.85-0.88 (0.87). Meta- tarsus I 0.10-0.11 (0.10), II 0.29-0.30 (0.30). DAGO 0.10, 0.10 n=2. n=3 except where shown.

Material examined: 1 ex., Bucks, Slough, -.VIII.61, Cerastium sp., GEW. 3 ex., Perths, Kinloch Rannoch, 26.VI.78, GMcG.

Remarks: Nymphs were found running on an upland heath dominated by Erica tetralix, Thymus drucei and - 3 1 3 -

Figure 69

•

Chlamydatus saltitans C. pullus

Strongylocoris leucocephaIus - 314 -

Galium saxatile, along with nymphs of Plagiognathus chrysanthemi to which they bear a superficialiesemblance. They can be separated by the fact that the nymphs of P. chrysanthemi are slightly larger, do not have red eyes and lack spots on the pronotal angles. Chlamydatus wilkinsoni (two adult d, one adult 0 was also found, but no nymphs were obtained. The adult of Pachytomella parallela,found in the same locality, were very similar to the adults of C. wilkinsoni, but could easily be distinguished by examination of the pretarsus. The former have flap-like parempodia (Orthotylinae) and the latter bristle-like parempodia (Phylinae). The metathora- cic femoro-tibial articulation in Chiamydatus appears more specialised and it is conceivable that some energy storage mechanism may be involved in its saltatorial ability. The metathoracic femora are greatly expanded. For further discussion see Halticus saltator.

See also Figures 14f; 15a,b C. wilkinsoni - 315 - Monosynamma bohemani (Fallen) Figs.15c,d; 19:56,57; 25:5; 54

Antennal segment I lightly fuscous, rest of antenna pale. Rostral segment IV fuscous. Abdomen paler than head, nota and legs, terminal segment fuscous, each segment with one faint spot on each lateral margin. Dorsal body hairs short- ish, black, simple and stout,long, black, bifurcate, occasionally trifurcate (max.). 0.10). Anterior margins of pro- and meso-thoracic femora with one stout spine distally, metathoracic femora with two spines. Tibial spines shorter than tibial diameter, arising from minute dark spots. DAGO broad, double with anterior bar sclerotised, sinuate, and surrounded by pigmented cuticle. Claws striate basally. Pulvilli medium. Parempodia long, bristle-like. Trichobothria 6:5, trichs long, trichomae compact. Bothria recessed.

Measurements: Total body 1. 2.69-3.20 (3.02). Head w. 0.72- 0.80 (0.75),1. 0.48-0.60 (0.56), vertex 0.35-0.43 (0.39). Antenna total 1. 1.66-1.76 (1.69) n=5, I 0.22-0.26 (0.24), II 0.60-0.64 (0.61), III 0.43-0.48 (0.45) n=6, IV 0.38-0.40 (0.39) n=5. Rostrum total 1. 1.16-1.22 (1.20), I 0.30,0.34 n=2, II 0.30-0.34 (0.32) n=3, III 0.22-0.24 (0.23) n=3, IV 0.30-0.32 (0.31) n=6. Pronotum a.w. 0.59-0.68 (0.67), p.w. 0.80-0.88 (0.84), 1. 0.38-0.42 (0.40). Wing pad 0.46-0.52 (0.49). Metafemur 1. 0.90-0.98 (0.94), w. 0.26-0.28 (0.27).= Metatibia 1. 1.24-1.35 (1.29). Metatarsus) 0.12-0.14 (0.13), II 0.38-0.40 (0.39). DAGO 0.14-0.16 (0.15). n=7 except where shown.

Material examined: 7ex., Lancs, Sandscale, 11.VII.54, Salix repens, TRES. 4ex., Kent, Dungeness, 8.VII.65, GEW. 2ex., Kent, Dungeness, 15.VII.67, GEW.

Remarks: No reference to the live colour of the nymphs of this species has been found, but the head and nota are possibly brown, and the abdomen orange-red with a green tinge. Butler (1923) recorded Salix lapponum as a host plant. The nymphs are very similar to those of Psallus. - 316 - Camyylomma verbasci (Meyer-Dtlr) Figs. 19 60,61; 25:6

Pale blue-green. Pronotum with two faint transverse green bands. Posterior pronotal angles rounded. Dorsal body hairs black, short, simple or long, bifurcate, trifurcate (max.l. 0.9). Ventral body hairs slender, simple. Metathoracic femora with two dark spots on distal anterior edge, each bearing a stout spine. Metathoracic tibiae with 11 dark spots, each bearing a stout spine just longer than the tibial diameter. Meso- and meta-thoracic tibiae with fewer spines. DAGO single, pale with anterior bar straight. Claws thin,striate basally. Pulvilli medium. Parempodia bristle- like. Trichobothria.7:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 2.12-2.40 (2.26) n=5. Head w. 0.59-0.64 (0.61) n=7, 1. 0.41-0.48 (0.44) n=7, vertex 0.30- 0.34 (0.32) n=7. Antenna total 1. 1.18-1.22 (1.20) n=5, I 0.16-0.18 (0.17) n=7, II 0.39-0.44 (0.42) n=7, III 0.34-0.36 (0.35) n=5, IV 0.24-0.27 (0.26) n=5. Rostrum total 1. 0.90- 0.99 (0.95) n=7, IV 0.28-0.30 (0.29) n=7. Pronotum a.w. 0.50-0.56 (0.52) n=6, p.w. 0.66-0.74 (0.70) n=6, 1. 0.28- 0.32 (0.30) n=6. Wing pad 0.38-0.40 (0.39) n=8. Metafemur 1. 0.57-0.70 (0.64) n=8., w. 0.20-0.24 (0.23) n=8. Meta- tibia 1. 0.82-0.97 (0.91) n=7. Metatarsus I 0.10-0.11 (0.10) n=7, II 0.30-0.32 (0.31) n=7. DAGO 0.037-0.046 (0.044) n=7.

Material examined: 7ex., Kent, Gravesend, 8.VIII.49, Verbascum thapsus, TRES.

Remarks: Southwood and Leston (1959) recorded this species as being an important predator of red spider mites, and aphids, as well as feeding on several plants, including hollyhock, mullein and . - 317 -

Salicarus roseri (Herrich-Schiffer) Figs. 19:62,63; 25:7

Posterior margins of nota fuscous. Dorsal body hairs dense, black, short, curved, parallel-sided, bifurcate, trifurcate or serrate. Latero-ventral body hairs as dorsal hairs, medio-ventral hairs slender, simple (max.l. 0.07). Meta- thoracic tibiae with 3 rows of 15 stout striate spines, not as long as tibial diameter. Meso- and pro-thoracic tibiae with fewer spines. Cuticular sculpturing dense as small pimples. DAGO single, anterior bar long,straight, sclerotised, margins dark, with small pigmented patch posteriad. Claws strong basally. Pulvilli medium. Parempodia bristle-like. Trichobothria 7:6, trichomae present on some. Bothria slightly recessed.

Measurements: Total body 1. 3.00-3.60 (3.40). Head w. 0.76- 0.82 (0.79), 1. 0.48-0.54 (0.52), vertex 0.40-0.48 (0.44). Antenna total 1. 1.26-1.38 (1.32), I 0.20-0.26 (0.23), II 0.48-0.54 (0.50), III 0.30-0.32 (0.31), IV 0.28-0.30 (0.29) Rostrum total 1. 1.12-1.20 (1.16), 10.30-0.36 (0.33), II 0.28-0.32 (0.30), III .0.22-0.26 (0.23), IV 0.28-0.32 (0.30). Pronotum a.w. 0.64-0.80 (0.72), p.w. 0.96-1.04 (1.00), 1. 0.44-0.48 (0.46). Wing pad 0.46-0.54 (0.52). Metafemur 1. 0.82-0.92 (0.87), w. 0.30-0.32 (0.31). Metatibia 1. 1.20- 1.32 (1.25). Metatarsus 10.12-0.14 (0.12), II 0.34-0.40 (0.36). DAGO 0.060-0.062 (0.061). n=10 in all cases.

Material examined: 10ex., Herts, Elstree, 20.VI.60, Salix sp., DL.

Remarks: The pubescence of the nymphs of this species is characteristic. Kullenberg (i944) sketched a nymph with a dark red abdomen, brown head and nota, and pale legs and antennae. The terminal abdominal segment is shown dark and there are faint lateral spots on each segment. - 318 -

Sthenarus rotermundi (Scholtz) Figs. 19:64,65; 25:8

Rostral segment IV, apical third of tarsal segment II fuscous. Dorsal body hairs stout, long, black, bifurcate or serrate (max.l. 0.14), and shorter, expanded apically, bifurcate, trifurcate or coronate. Ventral body hairs straight, simple. Rostrum, antennal segments III, IV, tarsi with simple hairs. Metathoracic femora with 3 spines on distal anterior margin. Meta- and meso-thoracic tibiae with stout, striate spines, longer than tibial diameter, arising from small fuscous spots. Prothoracic tibiae with short spines. Dorsal cuticu- larsculpturing immediately posterior to DAGO in polygonal pattern. DAGO broad, double, anterior bar sinuate, weakly sclerotised. Claws stout, striate. Pulvilli medium. Parem- podia bristle-like. Trichobothria 7:6, trichomae present on some. Bothria slightly recessed.

Measurements: Total body 1. 2.80-3.28 (3.06). Head w. 0.80- 0.84 (0.83),1. 0.55-0.62 (0.59), vertex 0.46-0.52 (0.49). Antenna total 1. 1.65-1.72 (1.69), I 0.24-0.27 (0.25 ), II 0.69-0.75 (0.71), III 0.39-0.42 (0.41), IV 0.32-0.36 (0.33). Rostrum total 1. 1.12-1.18 (1.16) n=9, IV 0.30-0.32 (0.31) . Pronotum a.w. 0.72-0.80 (0.77), p.w. 0.96-1.04 (1.01), 1. 0.44-0.50 (0.47). Wing pad 0.48-0.54 (0.52). Metafemur 1. 0.91-0.98 (0.96), w. 0.34-0.40 (0.36) . Metatibia 1. 1.28- 1.36 (1.32). Metatarsus I 0.12-0.16 (0.13) n=9, II 0.37- 0.40 (0.38),n=9. DAGO 0.12-0.14 (0.13). n=10 except where shown.

Material examined: 11ex., Berks, Silwood Park, 13.VI.57, TRES,

Remarks: The general colour of the nymphs is probably pale grey-green. Southwood and Leston (1959) recorded this species as being confined to white aid grey poplars. - 319 - Asciodema fieberi (Douglas and Scott) Figs. 26:1; 53

Dorsal body hairs white, long, bifurcate, trifurcate and smaller, simple. Tibial spines short, pale..DAGO pale, single with anterior bar curved, not very sinuate. Claws straight, curving apically. Pulvilli large, rugulose but not reaching claw apices. Parempodia short, bristle-like. Trichobothria 7:6, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 2.68, 2.76. Head w. 0.70, 0.70, vertex 0.36,0.40. Antenna segment II 0.69, 0.71, III 0.49, 0.52. Pronotum a.w. 0.55, 0.56, p.w. 0.78, 0.80. Wing pad 0.50, 0.52. Metatibia 1. 1.36, 1.40. Meta- tarsusl 0.13, 0.14, II 0.36, 0.39. DAGO 0.06, 0.06.

Material examined: 2 ex., Berks, Beenham Hill, -.VI.69, GEW.

Remarks: Butler (1923) described the nymphs ' delicate green with white legs aid antennae, wing pads yellowish, abdomen showing its dark contents through the skin, apex of rostrum black - 320 -

Asciodema obsoletum (Fieber) Figs. 19:66,67; 26:2; 42c

Greyish-green. Abdomen, tarsi and apex of rostrum with fus- cous tinge. Dorsal body hairs dense, pale, dark, long, bi- furcate or trifurcate (max.l. 0.11). Ventral body hairs simple. Meso- and metathoracic tibiae with stout spines just longer than tibial diameter. DAGO oval, single, pale with anterior bar straight. Claws curved, striate. Pulvilli medium. Parempodia bristle-like. Trichobothria 6:6, trichomae present on most. Bothria slightly recessed.

Measurements: Total body 1. 2.84-3.04 (2.98). Head w. 0.71- 0.74 (0.73), 1. 0.54-0.62 (0.58), vertex 0.40-0.44 (0.43). Antenna total 1. 1.86-2.03 (1.95) n=6, 10.27-0.28 (0.28), II 0.64-0.74 (0.69), III 0.55-0.62 (0.59), IV 0.36-0.42 (0.38) n=6. Rostrum total 1. 1.28-1.32 (1.31), IV 0.30-0.34 (0.32). Pronotum a.w. 0.64-0.67 (0.65), p.w. 0.80-0.86 (0.84), 1. 0.40-0.42 (0.40). Wing pad 0.46-0.52 (0.49). Metafemur 1. 0.94-1.00 (0.98), w. 0.30-0.34 (0.32). Metatibia 1. 1.38- 1.44 (1.42). Metatarsus 10.12-0.14 (0.13), II 0.39-0.43 (0.41). DAGO 0.060-0.067 (0.064). n=7 except where shown.

Material examined: 11 ex., Berks, Silwood Park, 200V1.50/ 23.VII056, Sarothamnus scoparius, TRES. 5 ex., Herts, Harpenden, 6.VII.54, Ulex europaeus, TRES.

Remarks: Waloff and Southwood (1960) described the nymphs of this species, discussed their biology and included them in a key to the immature stages of mirids occurring on broom. - 321 - Hallodapus montandoni (Reuter) Figs. 2d; 15e; 26:3

Red and white. Base, apex of antennal segment I, apical half of segment II, segment III and basal half of segment IV red. Head, nota, abdomen red to reddish-brown. Body with transverse, broad,white band on abdominal segments I, II and wing pads, extending to ventral surface including coxae. Pro- and meso-thoracic tibiae with proximal apices red, metathoracic tibiae with proximal half red. Tarsi slightly tinged brown. Rostrum long, reaching past meta- thoracic coxae. Pronotum constricted slightly anteriad. Dorsal body hairs long, silver with swollen or spatulate apices. Interommatidial setae simple. Metafemoral dorsal surfaces with areas of specialised cuticular sculpturing in form of patches of coarse ovoid bumps. DAGO oval, small, single with anterior bar short, straight, sclerotised. Claws straight. Pulvilli small. Parempodia bristle-like. Tricho- bothria 7:6, trichomae well-developed, compact. Bothria recessed.

Measurements: Total body 1. 3.00. Head w. 0.68, 1. 0.56, vertex 0.34. Antenna total 1. 2.52,1 0.38,I1 0.88, III 0.66, IV 0.60. Rostrum total 1. 1.40, IV 0.39. Pronotum a.w. 0.60, p.w. 0.80, 1. 0.44. Wing pad 0.48. Metafemur 1. 1.06, w. 0.22. Metatibia 1. 1.48. Metatarsus 10.17, II 0.42. DAGO 0.046.

Material examined: 3 ex., Kent, Eccles, 16.VII.60/1.VIII.60, AMM. 2 ex., Dorset, Cranborne, 20.VII.60, GEW.

Remarks: The metathoracic,femoral, dorsal sculpturing and the shape of the dorsal hairs are characters unique to the tribe Hallodapini. The wing pads are quite short, only reaching the posterior margin of the second abdominal seg- ment, this condition giving rise to bracypterous adults. Southwood and Leston (1959) recorded this species as preda- cious on the ant Myrmica scabrinodis. - 322 -

Hallodapus rufescens (Burmeister) Figs. 2e; 15f; 19:68,69; 26:4

Reddish-brown. Base of antennal segment I, apical half of segment II, segment III, basal half of segment IV red. Head, nota, abdomen reddish-brown. Abdominal segments I, II white, continuing to wing pads and extending to ventral surface of body. Femora red. Metathoracic tibiae with broad red bands in proximal half. Pronotum reaching to meso- thoracic coxae. Pronotum with lateral margins subparallel. Dorsal body hairs silver, long, swollen or spatulate apically (max.l. 0.15). Ventral body hairs long, simple. Inter- ommatidial setae simple. Cuticular sculpturing in four of large raised pimples or spots on head, nota, becoming more spinule-like on legs, abdomen, antennae. Metathoracic femora with areas of specialised cuticular sculpturing in form of patches of coarse, ovoid bumps. DAGO oval, single, with anterior bar short. Claws very straight. Pulvilli small. Parempodiabristle-like. Trichobothria 7:6, trichomae well-developed, compact. Bothria recessed.

Measurements: Total body 1. 3.04, 3.28. Head w. 0.68, 1. 0.60, vertex 0.36. Antennal segment I 0.40,11 0.96, 1.06, III 0.70, 0.76., Rostrum total 1. 1.27, 1.37, IV 0.34, 0.37. Pronotum a.w. 0.60, p.w. 0.76, 1. 0.40. Wing pad 0.50. Metafemur 1. 1.00, 1.20, w. 0.23,0.24. Metatibia 1. 1.44, 1.70. Metatarsus I 0.12;0.17, II 0.38, 0.48.

Material examined: 1 ex., Surrey, Oxshott, 19.VII.53, AMM. 2 ex., Surrey, Chobham, 26.VII.58, GEW. 1 ex., Surrey, Chobam, 21.VI.59, GEW. 1 ex., Isles of Scilly, Tresco, -.VI.59, GEW.

Remarks: The cuticular sculpturing on the metathoracic femora and the characteristic dorsal body hairs are similar to those found in the preceding species. The tibiae were not found to be particularly spotted, nor the tarsi black as described by Butler (1923), neither are the tibiae par- ticularly elongate. The lateral abdominal cuticular sculp- turing in the vicinity of the metathoracic femora changes - 323 - from the normal spinules to the raised ovoid bumps found on the metathoracic femora. This also occurs in the pre- ceding species, but the function is unknown (see discussion). - 324 - Systellonotus triguttatus (L.) Figs. 19:70,71; 26:5

Brownish-red. Antennal segment I pale brownish-red, rest of antenna brownish-red. Head, nota brownish-red with pinkish- red ecdysial lines. Abdomen brownish-red, segments I, II with broad,white, transverse band extending across wing pads to ventral body surface. Tibiae, femora brown, tibial apices pale brown. Tarsi fuscous. Head constricted posteriad. Pro- notum with lateral margins parallel. Abdomen swollen, rounded. Dorsal body hairs long, swollen or spatulate apically (max.l. 0.13). Interommatidial setae swollen or spatulate apically. DAGO oval, single, small , margins dark, anterior bar short, slightly curved. Claws quite straight. Pulvilli small. Parempodia bristle-like. Trichobothria 7:5, trichomae developed, compact. Bothria recessed.

Measurements: Total body 1. 3.55-3.88 (3.73). Head w. 0.72- 0.80 (0.75), 1. 0.70-0.74 (0.72), vertex 0.36-0.44 (0.39). Antenna total 1. 2.64-3.20 (2.99) n=3, I 0.38-0.40 (0.39), II 0.86-1.00 (0.94), III 0.86-1.00 (0.93) n=4, IV 0.48-0.80 (0.78) n=3. Rostrum total 1. 1.38-1.44 (1.43), IV 0.34- 0.39 (0.37). Pronotum a.w. 0.60-0.70 (0.67), p.w. 0.80- 0.81 (0.80), 1. 0.50-0.54 (0.52). Wing pad ð 0.52-0.54 (0.53) n=3. Metafemur 1. 1.08-1.24 (1.17), w. 0.20-0.25 (0.23). Metatibia 1. 1.62-1.80 (1.72). Metatarsus 10.14 0.18 (0.16), II 0.43-0.56 (0.49). DAGO 0.035-0.040 (0.038). n=5 except where shown.

Material examined: 6 ex., Bucks, Slough, -.VI.53, GEW. 4 ex., Bucks, Slough, -.VI.56 GEW. 2 ex., Kent, Eccles, 19.VIII.57, AMM. 2 ex., Surrey, Witley Common, -.VI.58, GEW.

Remarks: The nymphs of this antmimetic species have been described by Butler (1923) and sketched by Kullenberg (1944). nymph The ? Vth insta ha very short wing pads and the d long wing pads, reflecting the degree of development of adult wings. Butler (1923) associated the species with several ant species, fusca, Lasius niger and L. alienus. Southwood and Leston (195 9) recorded it as feeding on various plants, Calluna sp., Potentilla erecta and Salix repens, as well as - 325 - sucking ant pupae, aphids, but never live ants. This species shares the specialised dorsal hairs of the preced- ing two species, but lacks the cuticular sculpturing of the metathoracic femora. The spatulate hairs are very extensive in this species, even occurring between ouuuatidia on the tibiae and antennal segments (except segment IV). It is possible that there are two generations per year, for although the majority of nymphs have been recorded in June, two Vth instar nymphs were caught in mid-August, 1957. - 326 -

Dicyphus constrictus (i3oneman) (S. Dicyphus s . s . ) Figs. 19:72,73; 27:3; 55; 70

Pale green without distinct markings. Antennal segment I with broad,transverse, red band. Eyes dark red. Metatarsal segment II infuscate apically. Head tapers posteriad. Rost- rum reaches metathoracic coxae, segment IV stout basally, tapering to 0.25 basal width. Eyes remote from pronotum. Pronotum with anterior tranverse constriction. Metatarsal segment II 3.5x segment I. Dorsal body hairs long, pale, simple (max.l. 0.17). Ventral body hairs shorter,simple. Interommatidial setae sparse, simple. Metatibial spines pale (max.l. 0.016). Body cuticular sculpturing sparse. DAGO very small, single, pale,indistinct. Claws slender, striate, slightly recurved. Pulvilli large, flap-like, truncate, arising from basal claw projection. Parempodia bristle- like. Trichobothria 6:5, trichomae present, diffuse - femo- ral cuticle otherwise smooth. Bothria tuberculate.

Measurements: Total body 1. 3.32-3.56 (3.47)1-1=3. Head w. 0.60-0.62 (0.61) n=3, 1. 0.67-0.68 (0.68) n=3, vertex 0.25 -0.27 (0.26) n=3. Antenna 10.35-0.36 (0.36) n=3, II 0.78-0.82 (0.81) n=3. Rostrum total 1. 1.49-1.52 (1.50) n=2, IV 0.36-0.40 (0.39) n=3. Pronotum a.w. 0.54-0.62 (0.58) n=3, p.w. 0.60-0.67 (0.64) n=3, 1. 0.48-0.51 (0.49) n=3. Wing pad 0.50-0.61 (0.56) n=3. Metafemur 1. 1.27- 1.28 (1.28) n=3, w. 0.18-0.19 (0.18) n=3. Metatibia 1. 1.80-1.88 (1.85) n=3. Metatarsus I 0.16-0.18 (0.17) n=3, II 0.64-0.67 (0.65) n=3.

Material examined: 3 ex., Dumfrieshire, Bodesbeck Farm, 2.IX.76, Stachys sylvatica, CWM.

Remarks: This species, which was previously described by Butler (1923), is similar to D. stachydis, differing in having paler legs. The transverse red band on antenna seg- ment I is broader than in D. pallicornis. — 327 —

Figure 70

0.62 ' 060 Dicyphus constrictus (5th.) Dicyphus epilobli (5th.)

Dicyphus errans (5th.) 0.74 Dicyphus stachydis (5th.) - 328 -

Dicyphus epilobii Reuter (S. Dicyphus s.s.) Figs. 19:74,75; 27:4; 55; 70

Pale green. Antennae with segment I red; segment II with red tinges basally. Head with pale red stripe. Eyes red. Apices of rostral segment IV and tarsal segment II infuscate. Rostral segment IV tapering from 0.08 basally to 0.01 apically. Dorsal body hairs pale, simple (max.l. 0.21). Ventral hairs simple. Rostral IV hairs long, thin (max.l. 0.04). Posterior abdominal segment with dorsal cuticular sculpturing as small slightly curved groups of 6-8 spinules in transverse rows. DAGO small, oval, single, indistinct. Claws long, slender, recurved with thick basal projections. Pulvilli short, striate, flap-like. Parempodia bristle- like. Trichobothria 7:6, trichomae present, diffuse. Bothria tuberculate.

Measxrements: Total body 1. 3.72-4.64 (4.12) n=6. Head w. 0.62-0.70 (0.65) n=6, 1. 0.62-0.74 (0.66) n=6, vertex 0.20- 0.28 (0.26) n=6. Antenna total 1. 1.96-2.82 (2.35) n=5, I 0.32-0.42 (0.39) n=6, II 0.72-1.08 (0.96) n=6, III 0.52- 0.76 (0.70) n=5, IV 0.40-0.56 (0.45) n=5. Rostrum total 1. 1.40-1.70 (1.58) n=5, I 0.40-0.42 (0.40) n=5, II 0.36-0.48 (0.44) n=5, III 0.30-0.38 (0.34) n=5, IV 0.34-0.42 (0.40) n=5. Pronotum a.w. 0.48-0.60 (0.54) n=6, p.w. 0.64-0.88 (0.73) n=6, 1. 0.44-0.54 (0.50) n=6. Wing pad 0.58-0.68 (0.61) n=6. Metafemur 1. 1.16-1.60 (1.52) n=6, w. 0.18-0.24 (0.20) n=6. metatibia 1. 2.12-2.32 (2.11) n=5. Metatarsus 10.14-0.18 (0.16) n=6, II 0.54-0.68 (0.63) n=6.

Material examined: 3 ex., London, Peckham, 6.VII.48, TRES. 3 ex., Oxon, Little Wittenden Wood, 25.VI.50 TRES. 10 ex., Berks, Silwood Park, 3.VIII.77, Epilobium hirsutum, GMcG. 2 ex., Bucks, Slough, 16.IX.77, E. hirsutum, GMcG.

Remarks: Fifth instar nymphs of this species were collected at the Pest Infestation Laboratory (PIL) at Slough on 16.IX.77. This is quite late in the year and is further evidence that there is a second generation (Southwood & Leston,1959). The species was previously described by Butler (1923). - 329 -

Dicyphus errans (Wolff,J.F.) (S. Dicyphus s.s.) Figs. 19:76,77; 27:5; 43d; 55; 70; Pis 18; 19

Green to pale green. Antenna segment I red. Head and nota reddish green. Apical half of second tarsal segment and apical portion of rostral segment IV infuscate. Dorsal and ventral body hairs simple, long (max.l. 0.19). Posterior abdominal segment with cuticular sculpturing as small curved clumps of 6-8 spinules in transverse rows. DAGO small, single, with anterior bar very weak. Claws long, slender, with thick basal projections. Pulvilli flap-like. Parem- podia bristle-like. Trichobothria 6:5, trichomae present, diffuse. Bothria tuberculate.

Material examined: 1 ex., Bucks, Datchet, Lamiuor sp., TRES. 1 ex., Bucks, Denham, 29.VII.77, Stachys sylvatica, GMcG.

Remarks: The 'faint traces of spots' mentioned in the previous description by Butler (1923) are most probably the trichomae; otherwise there are no spots. The claws are stouter and less recurved than in D. epilobii. - 330 -

Plate 18 Dicyphus errans (Vth instar) x20 - 3 31 -

Plate 19 Dicyphus errans (Vth instar) x20 The nymph is cleaning its rostrum and antennae with the prothoracic tibial comb. - 332 -

Dicyphus stachydis Reuter (S. Dicyphus s.s.) Figs. 5f; 16a; 19:78,79; 27:6; 55; 70

Green. Antennd segment I with broad transverse reddish band. Head reddish-green. Apical portion of tarsal segment II and apex of rostral segment IV infuscate. Rostrum segment IV acutely tapered to apex. Dorsal body hairs long, pale, simple (max.l . 0.19) . Posterior abdominal segment with cuti- cular sculpturing as small clumps of 2-8 spinules in trans- verse rows. DAGO small, oval, single. Claws long, slender, slightly recurved, with large basal projections. Pulvilli flap-like. Parempodia bristle-like. Trichobothria 6:5, trichomae present, diffuse. Bothria tuberculate.

Measurements: Total body 1. 3.12-3.72 (3.32) n=10. Head w. 0.56-0.62 (0.60) n=10, 1. 0.56-0.62 (0.59) n=10, vertex 0.14- 0.20 (0.18) n=10. Antenna total 1. 2.20-2.54 (2.31) n=10, I 0.32-0.38 (0.33) n=10, II 0.76-0.94 (0.84) n=10, III 0.66- 0.76 (0.70) n=10, IV 0.42-0.48 (0.44) n=10. Rostrum total 1. 1.40-1.60 (1.51) n=10, I 0.36-0.40 (0.37) n=10, II 0.34-0.42 (0.39) n=10, III 0.30-0.38 (0.35) n=10, IV 0.40-0.42 (0.40) n=10. Pronotum a.w. 0.46-0.64 (0.53) n=9, p.w. 0.60-0.66 (0.62) n=9, 1. 0.46-0.58 (0.48) n=10. Wing pad 0.48-0.60 (0.54) n=10. Metafemur 1. 1.18-1.28 (1.23) n=10, w. 0.18- 0.22 (0.20) n=10. Metatibia 1. 1.74-1.92 (1.82) n=10. Metatarsus 10.16-0.20 (0.18) n=10, II 0.60-0.64 (0.62) n=10.

Material examined: 4 IVth instar ex., Herts, Harpenden, 23.VII.53, Stachys sylvatica, TRES. 11 ex., Hants, Alice Holt Forest, 25.VII.76, Lamium sp., JHM.

Remarks: This species is similar to D. errans and D. epilobii, but is distinguished by its smaller size, shorter legs, antennae and wing pads (Butler,1923). It was previously described by Butler (1923), who recorded Silene as the host for D. constrictus, Epilobium for D. epilobii, Lamium for D. errans and Stachys for D. stachydis. - 333 -

Dicyphus pallicornis (Meyer-Di1r) (S. Idolocoris) Figs. lc,d; 6a-d; 16b; 19:82,83; 27:7; 41c; 56; 71; P1 17

Green. Antennal segment I with red band near apex. Head pale. Eyes dark red. Rostral segment IV and apex of tarsal segment II slightly infuscate. Dorsal body hairs long, pale, simple (max.l . 0.14) . Ventral body hairs simple, shorter. Rostral segment IV tapered apically. Posterior abdominal segment with dorsal sculpturing as small clumps of 6-8 spinules in irregular rows. DAGO small, oval, single. Claws slender with basal projections. Pulvilli large, flap- like. Parempodia bristle-like. Trichobothria 6:4, trichomae present, diffuse. Bothria tuberculate. Proximal ventral metafemoral trichobothrium without trichomae.

Measurements: Total body 1. 3.12-3.44 (3.28) n=5. Head w. 0.60-0.66 (0.64) n=5, 1. 0.57-0.60 (0.59) n=5, vertex 0.25- 0.30 (0.27) n=5. Antenna total 1. 1.76-1.92 (1.88) n=5, I 0.26-0.30 (0.29) n=5, II 0.66-0.70 (0.70) n=5, III 0.48- 0.52 (0.50) n=5, IV 0.36 0.40 (0.39) n=5. Rostrum total 1. 1.22-1.44 (1.31) n=4, I 0.28-0.34 (0.31) n=3, II 0.29-0.32 (0.30) n=3, III 0.26-0.28 (0.27) n=4, IV 0.32-0.35 (0.34) n=5. Pronotum a.w. 0.56-0.66 (0.61) n=5, p.w. 0.76-0.78 (0.76) n=5, 1. 0.48-0.55 (0.51) n=5. Wing pad 0.46-0.52 (0.50) n=5. Metafemur 1. 1.04-1.14 (1.09) n=5, w. 0.20 (0.20) n=5. Metatibia 1. 1.38-1.50 (1.46) n=5. Metatarsus I 0.13-0.15 (0.14) n=5, II 0.48-0.50 (0.49) n=5.

Material examined: 3 ex., Isle of Rhum, Kinloch Wood, Digitalis sp., GEW. 8 ex., Berks, Silwood Park, 23.VI.77, Digitalis sp., GMcG. 12 ex., Notts, Radcliffe on Trent, 10 .VII.77 , (3 ex.),21.X.77 (2 ex.), 15.IV.78 (7 ex.), Digitalis sp., JHM.

Remarks: The nymphs which often have a dark green or black abdominal mass, are best collected by close examination of the basal rosettes of leaves and the stem. Sweeping is des- tructive and yields low numbers of nymphs. The numbers of Vth instar nymphs and adults found early in the season at Radcliffe on Trent suggests that these stages overwinter. The species was previously described by Butler (1923). - 334 -

Figure 71

0•56 058 ------Dieus pallicornis (5th.) ------Dicyphus annulatus (5th.)

0.64 0 82 Dicyphus rhododendri (5th.) bicyphus globulifer (5th.)

Campyloneura virgula (5th.)

3.4 - 335 -

Dicyphus annulatus (Wolff, J.F.) (S. Brachyceraea) Figs. 19:84,85; 27:8; 56; 71

Grey-green. Antennal segments with dark annulations, seg- ment IV. with red tinges. Head and pronotum dark with brown annulations. Abdomen dorsal surface with dark spots in transverse rows. Femorae with brown spots. Proximal and distal ends of tibiae, apical half of tarsal segment II and apical portion of rostrum segment IV with brown tinges. Rostrum reaching just past prothoracic coxae. Dorsal hairs simple (max.l. 0.13). Nota with dark hairs arisng from dark spots. Tibiae with dark spines arising from dark spots. Dorsal abdominal sculpturing as small slightly curved clumps of up to 8 spinules in transverse rows. DAGO small, oval, single with margins infuscate. Claws slender, slightly recurved with thick basal projections. Pulvilli large, flap- like. Parempodia bristle-like. Trichobothria 4:3, trichomae present, diffuse. Bothria tuberculate.

Measurements: Total body 1. 2.20-2.72 (2.39) n=10. Head w. 0.50-0.56 (0.52) n=10, 1. 0.44-0.52 (0.46) n=10, vertex 0.20- 0.24 (0.22) n=10. Antenna total 1. 1.12-1.24 (1.16) n=10, I 0.18-0.22 (0.20) n=10, II 0.40-0.42 (0.41) n=10, III 0.30- 0.34 (0.31) n=10, IV 0.22-0.26 (0.25) n=10. Rostrum total 1. 0.82-0.90 (0.85) n=10, I 0.20-0.24 (0.21) n=10, II 0.20- ,0.26 (0.22) n=10, III 0.20-0.24 (0.21) n=10, IV 0.20-0.24 (0.22) n=10. Pronotum a.w. 0.42-0.56 (0.45) n=10, p.w. 0.58-0.66 (0.61) n=10, 1. 0.32-0.38 (0.34) n=10. Wing pad 0.40-0.48 (0.43) n=10. Metafemur 1. 0.68-0.76 (0.71) n=10, w. 0.16-0.20 (0.18) n=10. Metatibia 1. 0.96-1.04 (1.00) n=10. Metatarsus I 0.10-0.12 (0.12) n=10, II 0.36-0.46 (0.38) n=10.

Material examined: 10 ex., Kent, Littlestone, 19.IX.55, Ononis sp., DL.

Remarks: This species was previously described by Butler (1923) . - 336 -

Dicyphus globulifer (Fallen) (S. Brachyceraea) Figs. 19:86,87; 28:1; 41a; 42f; 56; 71

Green. Antennal segment I with broad transverse dark band, base and apex of antennal segment II, apical half of segment III and whole of segment IV infuscate. Dorsal surface of head with dark markings. Apices of meso-and metanotal wing pads and apical half of tarsal segment II dark brown. Femorae with dark brown-black spots. Tibiae black at base and apex, with black spines. Antennal segment II with slight medial constriction. Rostrum reaching to prothoracic coxae. Dorsal surface hairs black, often arising from minute black spots. Posterior abdominal segments with cuticular sculp- turing as small compact clumps of 4-6 spinules in transverse rows. DAGO small, oval, single. Claws long, with large basal projections. Pulvilli large, flap-like. Parempodia bristle-like. Trichobothria 4:3, trichomae weak, diffuse. Bothria tuberculate.

Measurements: Total body 1. 3.16-3.68 (3.33) n=6. Head w. 0.60-0.64 (0.62) n=6, 1. 0.56-0.60 (0.59) n=6, vertex 0.22- 0.32 (0.26) n=6. Antenna total 1. 1.46-1.52 (1.49) n=6, I 0.24-0.28 (0.26) n=6, II 0.50-0.56 (0.53) n=6, III 0.40-0.42 (0.40) n=6, IV 0.28-0.30 (0.29) n=6. Rostrum total 1. 1.12- 1.14 (1.12) n=6, 10.30 (0.30) n=6, II 0.30 (0.30) n=6, III 0.22-0.24 (0.23) n=6, IV 0.28-0.30 (0.30) n=6. Pronotum a.w. 0.56 -0.62 (0.59) n=6, p.w. 0.44-0.46 (0.45) n=6, 1. 0.40- 0.56 (0.50) n=6. Wing pad 0.48-0.58 (0.55) n=6. Metafemur 1. 0.92-1.00 (0.96) n=6, w. 0.20-0.22 (0.21) n=6. Meta- tibia 1. 1.32-1.40 (1.35) n=6. Metatarsus I 0.14-0.16 (0.15) n=6, II 0.44-0.46 (0.45) n=6.

Material examined: 7 ex., Beds, Ampthill, 3.VIII.60, Silene alba, TRES. 6 ex., Bucks, Slough, 16.IX.77, GMcG.

Remarks: Six Vth instar nymphs and two IVth instar nymphs were collected at Slough 16.IX.77. The majority of this species become adult by late July, although nymphs have been recorded as late as early August. This species was described previously by Butler (1923). - 337 -

Dicyphus rhododendri Dolling,1972 Figs. 4f; 6e,f; 13; 16c; 19:80,81; 28:2; 55; 71

Red. Antennae pale ochreous, segment I and socket with dark markings, apex of segment and base of segment IV white. Head and nota with light brown markings. Eyes very dark, reddish- black. Legs pale ochreous, coxae white apically, apical half of tarsal segment II and whole of claws black. Head strongly tapered posteriad. Eyes remote from pronotum. Ros- trum reaching nearly to mesothoracic coxae. Dorsal body hairs simple (max.l. 0.12). Dorsal cuticular sculpturing on posterior abdominal segments as broken transverse rows of spinules or pimples. Femorae with cuticular sculpturing as fine spinules, elsewhere on body as thicker spinules or pimples. DAGO small, single, indistinct. Claws short, curved, without basal projections. Pulvilli lobe-like. Parempodia bristle-like. Trichobothria 3:3, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 3.24-3.82 (3.51) n=10. Head w. 0.52-0.62 (0.55) n=9, 1. 0.52-0.60 (0.56) n=10, vertex 0.20- 0.26 (0.23) n=9. Antenna total 1. 2.37-2.64 (2.52) n=10, 10.28-0.32 (0.30) n=10, II 0.72-0.78 (0.75) n=10, III 0.88- 1.00 (0.94) n=10, IV 0.49-0.60 (0.56) n=10. Rostra total 1. 1.10-1.19 (1.16) n=9, I (0.26) n=1, II (0.32) n=1, III (0.22) n=1, IV 0.28-0.31 (0.30) n=10. Pronotum a.w. 0.42-0.60 (0.52) n=10, p.w. 0.60-0.70 (0.65) n=10, 1. 0.40-0.48 (0.44) n=10. Wing pad 0.55-0.62 (0.60) n=10. Metafemur 1. 1.02-1.20 (1.12) n=10, w. 0.18-0.22 (0.20) n=10. Metatibia 1. 1.24-1.74 (1.63) n=10. Metatarsus 10.11-0.14 (0.12) n=105 II 0.37- 0.44 (0.40) n=10.

Material examined: 18 ex., Berks, Silwood Park, 3.VII.77, Rhododendron sp., GMcG.

Remarks: Second and third instar nymphs were collected on 23.VI.77 and IVth and Vth instar nymphs on 3.VII.77 at Silwood Park. This species, which was first described in Britain by Dolling (1972), seems to be confined to Rhododendron spp. and has_colonised most of the home counties since its capture at Kew, London, 1971. The species is - 338 -

predacious on aphids ( Masonaphis lambersi MacGillivray). The nymphs have not been described, but are easily distin- guished from the other Dicyphus species by being red,(the other species having green nymphs). The pretarsal char- acters, femoral sculpturing (absent in other Dicyphus sp.), the reduced trichobothrial number, the abdominal spiracular openings (much less expanded than in other Dicyphus sp.) and the colour of the nymphs point to the fact that this species is wrongly placed in Dicyphus. The British species of Dicyphus have long, slender claws which may be recurved and large basal projections, from which arise flap-like, truncate pulvilli. D. rhododendri has a short, curved claw, with no basal projection. Dolling finds that this species is probably closely related to the American 'Agilis- Cucurbitaceous' group, which is particularly nearctic. (pers. comm.); the nymphal characters agree with these find- ings. A new genus should be erected to contain D. rhododendri and its American relatives. Scanning electron microscopy clearly illustrates the difference in the pre- tarsal structures in this species as compared with other members of the genus (see figs. 6a-f). - 339 -

Campyloneura virgula (Herrich-Sch1ffer) Figs. 19:88,89; 28:3; 56; 71

Yellow. Antennal segment I, basal and medial portion of segment II, basal half of segment III and all of segment IV bright red in early instars; becoming less distinct in later instars, segment I and II, basal half of segment III and apex of segment IV red. Eyes red. Head with lateral mar- gins posterior to eyes with red stripe continuing down later- al margins of pro-, meso- and meta-nota . Wing pads largely red in later instars. Femora yellow, tinged red apically (the meso- and meta-thoracic tibiae of early instars are also tinged with red). Dorsal surface hairs simple. Dorsal surface with polygonal cuticular sculpturing (not as marked as in the Bryocorinae). Abdominal spiracles not expanded (diameter 0.008). DAGO double, pale, with anterior bar sinuate. Claws sub-apical, thick basally. Pulvilli large. Parempodia bristle-like. Trichobothria 8:6, trichomae absent (femoral spinules distributed evenly over entire surface). Bothria tuberculate.

Measurements: Total body 1.3.12-3.40 (3.31) n=7. Head w. 0.66 -0.68 (0.67) n=7, 1. 0.52-0.60 (0.57)n=7, vertex 0.28- 0.35 (0.32) n=7. Antenna total 1. 3.37-3.50 (3.44) n=5, I 0.36 -0.40 (0.38) n=7, II 1.32-1.36 (1.34) n=7, III 0.80- 0.84 (0.82) n=5, IV 0.86-0.92 (0.89) n=5. Rostrum total 1. 1.20-1.30 (1.24) n=7, rostral segment IV 0.28 (0.28) n=7. Pronotum a.w. 0.62-0.66 (0.64) n=6, p.w. 0.76-0.79 (0.77) n=7, 1. 0.40-0.42 (0.41) n=7. Wing pad 0.58-0.62 (0.61) n=7. Metafemur 1. 1.26-1.38 (1.31) n=7, w. 0.32-0.34 (0.33) n=7. Metatibia 1. 1.80-1.92 (1.85) n=7. Metatarsus I 0.14-0.17 (0.15) n=7, II 0.40-0.42 (0.41) n=7.

Material examined: 1 ex., Herts, Harpenden, 17.VIII.53, Acer pseudoplatanus,TRES. 2 ex., Herts, Harpenden, 16.VII.54, Acer pseudoplatanus,TRES. 7 ex., Herts, Borehamwood, 26.VI.66, Fraxinus sp., DL. 3 ex., London, Hampstead Heath, 28.VI.76,Crataegus sp., GMcG. 1 ex., Berks, Silwood Park, 16.VII.76, Rhododendron sp., GMcG. 2 ex., Exeter, North Exe, 4.VII.77, Scolopendrium vulgare, JGO - 340 -

Remarks: The nymphs of this distinctly marked species were previously described by Butler (1923). The species is predacious and has been recorded from a great variety of plants, including apple, cherry, ash, holly and a number of herbacious plants. The nymphs differ from all other Dicyphinae in having squat, subapical claws, non-expanded spiracles, and extensive polygonal sculpturing on the dorsal surface of the head, nota and Ist and Ilnd abdominal seg- ments. This sculpturing is a particularly bryocorine feature and it is interesting to observe that this species has recently been collected from Harts-tongue fern, Phyllitis scolopendrium. Campyloneura virgula is undoubtedly a parthenogenetic species, very few males ever having been found. All the nymphs examined were ?. It is possible that this species might be better placed in another group on the basis of its subapical claw and the other features peculiar to it, although it is not certain to which group it has the closest affinities. - 341 - Pilophorus cinnamopterus (Kirschbaum) Figs. 19:92,93; 28:4; P1 11

Reddish-yellow. Antennal segment I pale green, tinged red apically, segment II slightly incrassate apically with red tinge, segment III red, segment IV white, tinged with brown apically. Head and nota brown, tinged with red, antennal sockets black, ecdysial lines white. Eyes red, touching pro- notal margin. Pronotum with posterior margin white. Meso- and meta-nota with reddish-brown tinges, posterior margins black. Abdominal segment I with posterior half white, terminal segment fuscous, all segments with lateral fuscous patches ventrally. Margins of coxal joints black. Head wider than pronotum. Rostrum reaching just past mesothoracic coxae. Dorsal body hairs black, slightly curved, lanceolate. Tibial spines black, shorter than tibial breadth. DAGO broad, single, strongly sclerotised with anterior bar curved pos- teriad, pigmented patch anteriad and posteriad. Claws striate. Pulvilli small, adpressed. Parempodia flap-like, striate, truncate. Trichobothria 6:5, trichomae present, small, compact. Bothria recessed.

Measurements: Total body 1. 3.52-4.08 (3.76). Head w. 0.98- 1.04 (1.01), 1. 0.71-0.88 (0.79), vertex 0.44-0.52 (0.47). Antenna total 1. 2.56-2.80 (2.70), I 0.32-0.38 (0.35), II 1.18-1.32 (1.24), III 0.50-0.58 (0.55), IV 0.48-0.60 (0.55). Rostrum total 1. 1.37-1.61 (1.48), I 0.20-0.36 (0.29), II 0.34-0.39 (0.36), III 0.36-0.45 (0.38), IV 0.42-0.45 (0.44). Pronotum a.w. 0.84-0.89 (0.86) n=8, p.w. 1.00-1.08 (1.04) n=8, 1. 0.54-0.60 (0.58). Wing pad 0.52-0.64 (0.58). Meta- femur 1. 1.26-1.40 (1.34), w. 0.26-0.31 (0.28). Metatibia 1. 1.84-2.12 (2.03). Metatarsus 10.14-0.16 (0.15), II 0.41-0.45 (0.42). n=9 except where shown.

Material examined: 12 ex., Surrey, Longcross, 20.VII.77, Pinus sylvestris, GMcG. 6 ex., Berks, Silwood Park, 3.VIII.77, P. sylvestris, GMcG.

Remarks: The early instars of this predacious species are more strongly red. The nymphs were described by Butler (1923) and Kullenberg (1944) gave a colour drawing. - 342 - Pilophorus confusus (Kirschbaum)

Translated from Van Dinther (1953).

Dominant body colour blackish-brown. Antennal segment I dark grey; basal half of segment II brownish-yellow, apical half brownish-red; basal half of segment III whitish- grey, transparent, apical half brownish-red; basal half of segment IV white, transparent, apical half grey, tinged with brownish-yellow. Head blackish-brown, genae very light brown. Eyes reddish-brown. Pronotum brown with posterior margin white. Meso- and meta-nota with margins of wing pads blackish-brown. Abdomen blackish-brown with posterior margin of first segment white. Legs blackish-brown to brownish-yellow. Tarsi black. Dorsal surface of head, thorax, wing pads and abdomen with many toothed setae ( bifurcate, trifurcate, or coronate hairs - GMcG). Antennae hairy, especially the apical portion of segment IIA segment I with 3 or 4 well-developed setae with toothed apices.

Measurements: Total body 1. 3.60, breadth 1.25. Antenna total 1. 2.79, 10.31, II 1.22, III 0.56, IV 0.70.

Material examined: no nymphal material has been examined.

Remarks: This species has been recorded in Britain from low herbage and dwarf willows (Southwood & Leston, 1959). - 343 - Pilophorus perplexus Douglas and Scott Figs. 19:94,95; 28:7; 41f; 43a; P1 10

Antennal segment I light yellowish-brown; II similarly coloured basally, becoming strongly red apically; III red; IV with basal 3 white, apical 3 red. Head and nota brown,tinged with brownish-red, ecdysial lines brownish- yellow. Eyes dark red. Pronotum with posterior margin pale ochreous. Mesonotum with one dark spot on either side of ecdysial line. Abdomen reddish-yellow, segment I brown anteriad, white posteriad; II brown anteriad, reddish- yellow posteriad; terminal two segments brown. Pro- and meta-thoracic coxae white, mesothoracic coxae reddish- brown with brown transverse band. Femora and tibiae reddish- brown to brown. Tarsi brown. Rostrum reaching just beyond metathoracic coxae. Eyes touching pronotum. Pronotum convex with anterior angles rounded. Dorsal body hairs black, slightly curved, lanceolate. Ventral hairs simple. DAGO oval, single, sclerotised with anterior bar slightly curved posteriad, with pigmented patch anteriad and posteriad. Claws slightly curved, striate. Pulvilli very small. parempodia flap-like, striate. Trichobothria 6:5, trichomae present, compact. Bothria recessed.

Measurements: Total body 1. 3.68-3.88 (3.78) n=7. Head w. 0.92-1.02 (0.95) n=6, 1. 0.72-0.80 (0.77) n=6, vertex 0.40-0.50 (0.44) n=6. Antenna total 1. 2.50-2.69 (2.61) n=6, I 0.31-0.34 (0.32) n=7, II 1.12-1.20 (1.16) n=7, III 0.56 -0.58 (0.57) n=7, IV 0.48-0.59 (0.55) n=6. Rostrum total 1. 1.39-1.45 (1.42) n=5, I 0.30-0.34 (0.32) n=5, II 0.30-0.34 (0.32) n=5, III 0.37-0.38 (0.38) n=6, IV 0.38- 0.40 (0.40) n=6. Pronotum a.w. 0.76 n=1, p.w. 1.00 n=2, 1. 0.57-0.60 (0.60) n=5. Wing pad 0.41-0.56 (0.51) n=7. Metafemur 1. 1.14-1.32 (1.23) n=7, w. 0.26-0.30 (0.28) n=7. Metatibia 1. 1.86-1.98 (1.91) n=7. Meta-tarsus I 0.14- 0.16 (0.15) n=5, II 0.40-0.46 (0.43) n=5.

Material examined: 1 ex., Herts, St Albans, 26.V.53, Malus sp., TRES. 1 ex., Bucks, Slough, 2.VIII.64, TGEW. 18 ex., London, Putney Heath, 21.VII.77, Quercus sp., GMcG. 1 ex., Berks, Silwood Park, 3.VIII.77, Quercus sp., GMcG. - 344 -

Remarks: Younger instars show more red colouring. This predacimus species is found on a variety of deciduous trees (Southwood & Leston, 1959). - 345 - Halticus apterus (L.) Fig. 29:1

Brownish-black. Antennae pale ochreous, segment I slightly brown basally, IV very long with slight light brown tinge. Head and nota brownish-black with ecdysial lines pale ochreous. Eyes pale ochreous. Meso- and pro-thoracic legs pale ochreous. Metathoracic femora dark brown, shiny with white patch distally, extending on to tibiae. Metathoracic tibiae with proximal 3 (excluding the white patch) concolour- ous with femora, distal 3 pale ochreous. Claws and apical portion of tarsal segment II brown. Body small, squat. Meta- thoracic femur greatly expanded. Dorsal body hairs black, stout, long, simple, some arising from minute black spots. DAGO single, anterior bar sinuate, pigmented. Claws striate, thick basally. Pulvilli small, pointed. Parempodia striate, flap-like. Trichobothria 7:6, trichomae developed. Bothria well-recessed.

Measurements: Total body 1. 3.16,3.28 (3.22). Head w. 0.80; 0.82 (0.81), 1. 0.80,0.84 (0.82), vertex 0.40,0.42 (0.41). Antenna total 1. 3.80, 3.85 (3.83), 0.33, 0.36 (0.35), II 1.22, 1.24 (1.23), III 0.96, 0.98 (0.97), IV 1.26,1.30 (1.28). Rostrum total 1. 0.85, 0.88 (0.87), I 0.26, 0.28 (0.27), II 0.26, 0.28 (0.27), III 0.12, 0.13 (0.13), IV 0.20, 0.20 (0.20). Pronotum a.w. 0.56, 0.61 (0.59), p.w. 0.76, 0.82 (0.79), 1. 0.40, 0.42 (0.41). Wing pad 0.46, 0.50 (0.48). Metafemur 1. 1.22, 1.24 (1.23), w. 0.36, 0.39 (0.38). Metatibia 1. 1.86, 1.94 (1.90). Metatarsus 10.14, 0.16 (0.15), II 0.44, 0.46 (0.45). n=2 for all measurements.

Material examined: 2 ex., Hants, Pamber Forest, 19.VII.65, GEW.

Remarks: The small, squat shape of the nymphs of this genus, their greatly expanded metathoracic femora and their very long antennae make them readily recognisable. Butler (1923) described the nymphs and Southwood and Leston (1959) gave Galium sp. as the common host plant. - 346 - Halticus saltator (Geoffroy) Figs. 16d-f; 19:96,97

Head and nota brown, shiny. Metathoracic femora brown. Claws on apical 3 of tarsal segment infuscate. Rest of nymph pale ochreous. Head broad. Eyes touching pronotum. Pronotum with anterior angles rounded. Metathoracic femora greatly expanded, with two spines on anterior margins. Dorsal body hairs long, pale, simple. Metathoracic tibia with some pale spines on distal half. DAGO single, anterior bar sinuate with pigmented patch anteriad and posteriad. Claws striate basally. Pulvilli small, adpressed. Parem- podia flap-like. Trichobothria very long, 7:6, trichomae very marked. Bothria well-recessed.

Measurements: Total body 1. 2.48, 2.80 (2.64) n=2. Head w. 0.76, 0.78 (0.77) n=2, 1. 0.50, vertex 0.46, 0.51 (0.48) n=2. Antenna segment I 0.22, II 0.78. Rostrum total 1. 1.10, III 0.12, 0.14 (0.13) n=2, IV 0.20, 0.20 (0.20) n=2. Pronotum p.w. 0.82, 0.85 (0.84) n=2, 1. 0.36, 0.36 (0.36) n=2. Wing pad 0.40, 0.44 (0.42) n=2. Metafemur 1. 1.00, 1.02 (1.01) n=2, w. 0.40, 0.42 (0.41) n=2. Metatibia 1. 1.42, 1.52 (1.47) n=2. Metatarsus I 0.12, II 0.32.

Material examined: 2 ex., Hants, Winchester, -.VII.25, EAB.

Remarks: H. saltator differs from H. apterus in not having dark colouration on the metathoracic tibiae and in having shorter antennae. There is a strong indication that the nymphs and adults of this and the preceding species possess an energy storage mechanism, possibly in the metathoracic tibial-femoral articulation, related to their saltatorial habit. Such systems have been described in other insects (Bennet-Clark, 1975,1976). In the locust, Schistocerca gregaria, a structure known as the semi-lunar process (SLP) in the metathoracic tibial-femoral articula- tion acts as an energy store. In the Vth instar nymphs of H. saltator and H. apterus there appears to be a structure of similar shape and in a similar position to the SLP in locusts. The nymphs have been described by Butler(1923). - 347 - Strongylocoris leucocephalus (L.) Figs. 57; 69

Yellowish-brown or yellowish-orange. Antennal segment I yellowish-brown, II, III and IV lightly infuscate. Eyes reddish-brown. Posterior portion and margins of wing pads infuscate. Abdomen often mottled brown. Tibiae and femora pale yellowish-brown with stout black spines. Tarsi and claws blackish-brown. Body broad, very convex, rounded. Head flattened, very broad, with eyes overlapping pronotum. Pronotum with anterior and posterior angles very rounded. Rostrum short, reaching prothoracic coxae only. DAGO minute, pale. Claws short, squat, thick basally. Pulvilli very small. Parempodia large, flap-like . Trichobothria 3:2, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 3.24-3.48 (3.37) n=3. Head w. 1.08 (1e08) n=3, 1. 0.54-0.62 (0.57) n=3, vertex 0.64-0.68 (0.66) n=3. Antenna total 1. 1.34, 1.42 (1.38) n=2, I 0.24-0.26 (0.25) n=3, II 0.42-0.50 (0.47) n=3, III 0.34- 0.36 (0.35) n=3, IV 0.32, 0.32 (0.32) n=2. Rostrum total 1. 0.88-0.91 (0.89) n=3, I 0.30-0.32 (0.31) n=3, II 0.24- 0.26 (0.25) n=3, III 0.15-0.16 (0.16) n=3, IV 0.18 (0.18) n=3. Pronotum a.w. 0.76, 0.80 (0.78) n=2, p.w. 1.20-1.28 (1.25) n=3,1. 0.54 (0.54) n=3. Wing pad 0.36-0.50 (0.44) n=3. Metafemur 1. 0.82-0.88 (0.85) n=3, w. 0.25-0.28 (0.27) n=3. Metatibia 1. 1.14-1.18 (1.16) n=3. Metatarsus I 0.12- 0.14 (0.13) n=3, II 0.29-0.30 (0.30) n=3.

Material examined: 1 exo, Surrey, Boxhill, 31.V.19, EAB. 2 ex., Kent, Boxley, 9.VI.56, AMM. 3 ex., Oxon, Baldhill, 16.VII.62, Galium sp., GEW.

Remarks: The shape of this nymph, previously described by Butler (1923),is unique to the genus. It is also unusual in having a very small DAGO and a greatly reduced tricho- bothrial number. The trichobothria occur at the coxal end of the femur, the metathoracic femur appearing to lack the normally stable ventral pair of trichobothria (nos 2 and 3). This species has been associated with Helianthemum vulgare, Campanula rotundifolia, Galium veruri (Southwood & Leston,1959). - 348 -

Strongylocoris luridus(Fallen) Figs. 19:98,99; 29:6

Body broad, very convex, rounded. Head broad, fl attened. Eyes overlapping anterior margin of pronotum. Pronotum broad with anterior and posterior angles very rounded. Rostrum reaching to prothoracic coxae or just beyond. Dor- sal body hairs long, simple. DAGO minute, pale. Claws thick basally. Pulvilli small. Parempodia flap-like, striate. Trichobothria 4:3, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 3.00, 3.20 (3.10). Head w. 1.04, 1.08 (1.06), 1. 0.60, 0.64 (0.62), vertex 0.60, 0.60 (0.60). Antenna total 1. 1.34, 1.42 (1.38), I 0.20, 0.26 (0.23), II 0.46, 0.48 (0.47), III 0.34, 0.34 (0.34), IV 0.34, 0.34 (0.34). Rostrum total 1. 1.10, 1.13 (1.12), I 0.38, 0.39 (0.39), II 0.30, 0.31 (0.31), III 0.18, 0.19 (0.19), IV 0.24, 0.24 (0.24). Pronotum a.w. 0.70, 0.80 (0.75), p.w. 1.18, 1.20 (1.19), 1.0.45, 0.50 (0.47). Wing pad 0.34, 0.35 (0.35). Metafemur 1. 0.85, 0.92 (0.88), w. 0.28, 0.28 (0.28). Meta- tibia 1. 1.20, 1.24 (1.22). Metatarsus I 0.13, 0.14 (0.14), II 0.32, 0.33 (0.33). n=2 for all measurements.

Material examined: 2 ex., Cornwall, Pentire Head, 13.VI.56, Jasione montana, TRES.

Remarks: This species is very similar to the preceding species in that they share an unusual body form, minute DAGO and modified trichobothrial pattern. - 349 - Pachytomella parallela (Meyer-Dilar) Figs. 19:100,101; 29:7; 57

Brown to brownish-black. Antennae pale brown, segment II incrassate, III tinged red. Head and nota brown to brownish-black, slightly shiny. Eyes white touching pro- notum. Abdomen brownish-black with posterior two segments black, all segments with one median and two lateral fuscous spots. Pro- and mesothoracic legs pale ochreous. Meta- thoracic legs concolourous with body, metathoracic tibiae black with long black spines. Tarsi and claws brownish- black. Pronotum with anterior angles rounded. Wing pads in d reaching nearly to Vth abdominal segment. Dorsal body hairs dense, brown or black, simple. DAGO oval, single, heavily pigmented with anterior bar large, curved, raised with pigmented patch anteriad and posteriad. Claws fairly slender, curved apically. Pulvilli small. Parempodia flap- like, with long bristle arising from below. Trichobothria 6:5, trichs very long, trichomae strong, compact. Bothria well-recessed.

Measurements: Total body 1. 2.32-2.48 (2.41). Head w. 0.81- 0.88 (0.84), 1. 0.42-0.46 (0.45), vertex 0.42-0.49 (0.45). Antenna total 1. 1.50-1.58 (1.53), I 0.24-0.26 (0.24), II 0.48-0.52 (0.50), III 0.46-0.50 (0.48), IV 0.30-0.32 (0.31). Rostrum total 1. 0.88-0.95 (0.91), I 0.28-0.30 (0.29), II 0.26-0.30 (0.28), III 0.14 -0.15 (0.14), IV 0.18-0.20 (0.19). Pronotum a.w. 0.60-0.68 (0.67), p.w. 0.77-0.82 (0.80), 1. 0.34-0.36 (0.35). Wing pad d 0.58-0.62 (0.60). Metafemur 1. 0.81-0.84 (0.83), w. 0.20-0.24 (0.22). Meta- tibial 1. 1.02-1.08 (1.04). Metatarsus 10.10-0.12 (0.110, II 0.30-0.33 (0.31). DAGO 0.08-0.10 (0.09). n=4 for all measurements.

Material examined: 2 ex., Perths, Glen Sherrup, -.VII.37, ARW. '2 ex., Perths, Trossachs, 23.VI.57, ARW. 1 ex., Dumfs, Bran Law, 28.VII.63, ECP-C. 7 ex., Perths, Rannoch School, 24,25.VI.78, Galium saxatile, GMcG. 26 ex., Perths, Kinloch Rannoch, 26.VI.78, Galium sp., GMcG. 7 ex., Dumfs, Bodesbeck Law, 8.VII.78, Galium saxatile, GMcG & MMM. - 350 -

Remarks: The nymphs of this species live very close to the ground among Galium saxatile and other low plants, and have been recorded from the less exposed sides of mountains. The adults may be swept but more will be collected by search- ing on the ground. There is strong sexual dimorphism in the degree of wing development. The adult ?? are brachy- pterous and have a broad abdomen, whereas the d3' are macropterous and are more slender. This is reflected by the Vth instar nymphs - the wing pads in the are blunt and short, just reaching the DAGO and those of the. d' are long, nearly reaching the Vth abdominal segment. The adult ? may be confused in the field with the slightly smaller adult ? of Chlamydatus wilkinsoni (Douglas and Scott). The black metathoracic tibiae of P. parallela and its pretarsal structures will distinguish them.

There is a strong similarity between the nymphs of this species and those of the closely related species Orthocephalus saltator, although the former are much smaller. There is little difference in the ratios of body measurements, overall size being the only difference. It is interesting to speculate on the evolution of this species, which appears to have colonised upland meadows and heaths, occurring very low to the ground. O.saltator on the other hand is found more in lowland meadows and commonly on taller plants. The trichobothria in the two species are about the same absolute size, but relative to the femur, much longer in P. parallela. - 351 -

Orthocephalus saltator (Hahn) Figs. 19:102,103; 29:8

Head and nota blackish-brown. Abdomen brownish-green with dark patches in midline of each segment, smaller patches laterally. Claws and tarsi black. Antennal segment II slightly swollen in apical half. Dorsal body hairs black, single, quite long on pronotum. Tibiae and femora with stout black spines. DAGO single, highly sclerotised with anterior bar thick, curved. Claws thick basally. Parempodia flap- like. Pulvilli small, adpressed. Trichobothria 8:6, trichs long, trichomae weak. Bothria recessed.

Measurements: Total body 1. 3.44-3.76 (3.54) . Head w. 1.06- 1.20 (1.10), 1. 0.70-0.80 (0.74), vertex 0.52-0.68 (0.59). Antenna total 1. 2.26-2.50 (2.36), I 0.30-0.36 (0.35), II 0.76-0.88 (0.82), III 0.66-0.76 (0.71), IV 0.44-0.52 (0.49). Rostrum total 1. 1.18-1.34 (1.23) n=4, III 0.18- 0.20 (0.19), IV 0.24-0.27 (0.26). Pronotum a.w. 0.80-1.00 (0.90), p.w. 1.02-1.28 (1.14), 1. 0.50-0.58 (0.54). Wing pad ? 0.20-0.42 (0.31)n=2, 6 0.70-0.72 (0.71) n=3. Metafemur 1. 1.34-1.48 (1.39), w. 0.41-0.52 (0.45). Metatibia 1. 1.76- 1.98 (1.85). Metatarsus I 0.18-0.19 (0.18), II 0.50-0.54 (0.52). DAGO 0.088-0.113 (0.092). n=5 except where shown.

Material examined: 5 ex., Bucks, Slough, -.VI.53, GEW. 1 ex., Oxon, Aston Rowant, 1.VII.65, GEW.

Remarks: Butler (1923) described the nymphs of this species and those of O. coriaceus (Fabricius). Of the latter he said ' entirely shining black, broader and more hairy than 0. saltator, centre of abdomen slightly tinged with red; antennae and legs stouter, femoral hairs longer and much coarser; tibial hairs more numerous and very closely set; whole insect rather larger; outer limb of wing pad very slightly tinged with brown. Last instar 3zitnit.' 0. saltator is associated with various Compositae and is found in neglected pastures etc. (Southwood & Leston, 1959). The wing pad of the Vth instar d extends well past the DAGO, those of the ? nymphs reaching only to the DAGO. - 352 -

Malacocoris chlorizans (Panzer) Figs. 17a; 19:104,105; 30:1; 59

Pale green. Antennal segments and legs slender and very pale green. Head rather pentagonal, tapering posteriad. Eyes re- mote from pronotum. Pronotum with anterior angles slightly rounded. Dorsal surface hairs long, silver, simple, bifur- cate and trifurcate (max.1. 0.08). DAGO small, indistinct, with small anterior orange patch (the gland sac showing through the cuticle). Claws slender, striate, thin, claw angle approaching 90°. Pulvilli quite large, reaching mid- way to claw apices. Parempodia flap-like. Trichobothria 8:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 2.56-2.96 (2.73) n=5. Head w. 0.53-0.58 (0.55) n=5, 1. 0.42-0.56 (0.49) n=5, vertex 0.30- 0.36 (0.34) n=5. Antenna total 1. 2.92-3.22 (3.02) n=3, I 0.38-0.40 (0.39) n=5, II 1.05-1.18 (1.10) n=4, III 0.76- 0.90 (0.83) n=4, IV 0.62-0.74 (0.70) n=3. Rostrum total 1. 1.28 (1.26) n=2, I 0.21-0.23 (0.22) n=2, II 0.27-0.29 (0.28) n=2, III 0.37 (0.37) n=2, IV 0.39-0.43 (0.41) n=2. Pro- notum a.w. 0.34-0.42 (0.39) n=5, p.w. 0.56-0.64 (0.60) n=5, 1. 0.29-0.34 (0.32) n=5. Wing pad 0.50-0.61 (0.55) n=5. Metafemur 1. 1.16-1.19 (1.17) n=2, w. 0.23(0.23) n=2. Meta- tibia 1. 1.84-2.00 (1.92) n=5. Metatarsus I 0.10-0.11 (0.11) n=5, II 0.36-0.39 (0.37) n=5. DAGO 0.035 (0.035) n=2).

Material examined: 2 ex., Bristol, Longashton, 31.VII.73, Malus sp., DMG. 1 ex., Bucks, Denham, 27.VII.76, Corylus avellana, GMcG.

Remarks: The head shape and the colouration ( which is very soluble in alcohol) of this species is distinctive. There are no dark or fuscous markings on the body, neither are there traces of the black markings on antennal segment I and the base of segment II, such as are present in the adult stages. The nymphs were previously described by Butler (1923). - 353 - Cyllecoris histrionicus (L.) Figs. 19:106,107; 30:4; 42b; 72

Pale greenish-grey with dark markings. Antennal segments I, II, apical half of segment III and entire segment IV reddish- brown to dark reddish-brown. Head with ventral reddish- brown markings, antennal socket reddish-brown. Eyes red to reddish-brown, colour continuing down lateral margins of head and pronotum. Mesothoracic wing pads with light brown markings anteriad. Posterior 3 abdominal segments with a pair of distinctive reddish-brown patches, one on either side of the body midline. Femora with longitudinal brown stripe on anterior margins and occasionally on posterior margins. Tibiae with fuscous patch at proximal apices. Tarsi infuscate with segment II slightly incrassate. Antennal segment I and apical half of segment II slightly incrassate. Rostral tip infuscate, reaching mesothoracic coxae. Pronotum with lateral margins flared posteriad. Wing pads reaching nearly to IVth abdominal segment. Dorsal body hairs simple, occasionally bifurcate on antennal segment I and pronotum. DAGO single, simple with margins sclerotised. Claws thick basally, claw angle less than 90°. Pulvilli quite large, striate. Parem- podia large, flap-like, truncate apically. Trichobothria 4:4 trichs short, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 4.74-5.40 (5.07). Head w. 0.82- 0.90 (0.85), 1. 0.74-0.83 (0.79), vertex 0.52-0.58 (0.55). Antenna total 1. 3.70-3.94 (3.76), I 0.74-0.80 (0.77), II 1.50-1.60 (1.55), III 0.94-1.04 (0.99), IV 0.46-0.50 (0.47). Rostrum total 1. 1.40-1.49 (1.44) n=6, 10.35-0.40 (0.38) n=4, II 0.37-0.39 (0.38) n=4, III 0.30-0.34 (0.32) n=6, IV 0.36- 0.40 (0.38) n=7). Pronotum a.w. 0.70-0.80 (0.75), p.w. 1.04- 1.16 (1.09), 1. 0.54-0.64 (0,58). Wing pad 0.79-0.92 (0.86) . Metaf emur 1. 1.40-1.66 (1.55), w. 0.25-0.28 (0.27) . Meta- tibia 1. 2.20-2.52 (2.37). Metatarsus 10.13-0.15 (0.14), II 0.38-0.42 (0.40). n=10 except where shown.

Material examined: 32 ex., Herts, Harpenden, 31.V.54 - 11.VI.54, Quercus sp., TRES. 3 ex., Berks, Windsor, 21.VI.61, GEW. 12 ex., Hants, New Forest (Witley Wood), 4.VI.76, Betula and Quercus spp., GMcG. 3 ex., Berks, Silwood Park, 17.VI.77, - 354 -

Figure 72

Cyllecoris histrionicus -(Vth instar nymph)- • - 355 -

Quercus sp., GMcG. 1 ex., Berks, Silwood Park, 23.VI.77, Quercus sp., GMcG.

Remarks: Mason (in Butler, 1923) considered the adult stages to be mimics of of the genus . Indeed the adults look superficially similar to Malthodes marginatus (Latr.) and Malthinus flaviolus (Pay.), but it is not known why they should mimic these beetles. The nymphs, which have been described by Butler (1923) and illustrated by Kullenberg (1944), are readily identifiable by their colouration and markings. - 356 -

Dryophilocoris flavoquadrimaculatus (DeGeer) Figs. 19:108,109; 30:3; 59; 73; P1 6

Green with reddish-brown and white markings. Antennal seg- ment I reddish-brown with black markings basally, segment II black basally, becoming pale ochreous to reddish-brown apically, segment III with apical reddish-brown, segment IV entirely light brown. Antennal sockets black. Eyes ringed with white, black marks posteriad continuing to anterior angles of pronotum. Pronotum with posterior half pale ochreous. Meso- and meta-nota with large pale ochreous triangular patches pointing towards the body midline. Abdomen green with medial white patches, segment III with posterior margin white, segment IV with anterior margin white. Femora with distal halves light reddish-brown, basal halves pale ochreous. Tarsi, claws and apex of rostral segment IV blackish-brown. Pronotum with posterior half expanded laterally. Dorsal body hairs short, squat, black, coronate, becoming slightly longer on posterior portion of abdomen. Cuticular sculpturing as minute raised spots or pimples. DAGO small, oval, single, with anterior bar straight. Claws short, striate, thick basally. Pulvilli medium, rugulose. Parempodia thick, flap- like, with apices truncate. Trichobothria 5:4, trichs short, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 4.12-4.80 (4.54) n=7. Head w. 0.84-0.90 (0.87) n=7, 1. 0.60-0.68 (0.64) n=8, vertex 0.50-' 0.58 (0.54) n=7. Antenna total 1. 2.67-2.85 (2.78) n=6, I 0.48-0.50 (0.48) n=8, II 1.08-1.20 (1.15) n=8, III 0.74- 0.80 (0.78) n=7, IV 0.34-0.38 (0.36) n=6. Rostrum total 1. 1.10-1.17 (1.13) n=4, I 0.30-0.33 (0.32) n=5, II 0.26-0.28 (0.27) n=3, III 0.24-0.28 (0.26) n=4, IV 0.30 (0.30)n=7. Pronotum a.w. 0.82-0.91 (0.85) n=8, p.w. 1.16-1.24 (1.22) n=8, 1. 0.58-0.64 (0.60) n=8. Wing pad 0.72-0.86 (0.79) n=8. Metafemur 1. 1.21-1.28 (1.25) n=7, w. 0.23-0.28 (0.25) n=7. Metatibia 1. 1.88-1.94 (1.92) n=7. Metatarsus I 1.13- 1.14 (1.14) n=7, II 0.34-0.36 (0.35) n=7. DAGO 0.04-0.05 (0.045) n=7.

Material examined: 3 ex., Herts, Harpenden, 21.V.53, Quercus sp., TRES. 3 ex., Herts, Harpenden, 1,6.V.54, Quercus sp., - 357 -

Figure 73

Dryophilocoris flavoquadrimaculatus (Vth instar nymph) - 358 - TRES. 1 ex., Bucks, Slough, 12.V.61, Quercus sp., GEW. 1 ex., Berks, Windsor, 21.V.61, Quercus sp., GEW. 1 ex., Berks, Silwood Park, 17.VI.77, Quercus sp., GMcG.

Remarks: The pattern of markings, the pronotal shape and the squat, black coronate hairs are distinctive features of the nymphs of this species, which were previously described by Butler (1923). One specimen was found to be parasitised by a braconid wasp, probably Leiophron sp. - 359 -

Globiceps flavomaculatus (Fabricius) Figs. 19:110,111; 30:5

Antennae blackish-brown with segment I reddish-brown. Head blackish-brown. Pronotum blackish-brown with posterior angles tinged white. Abdomen green with red tinges, segment III tinged white; posterior two segments blackish-brown; other segments with central dark spot and two lateral spots each. Femora and tibiae reddish-brown. Tarsi blackish- brown. Head tapering posteriad. Dorsal body hairs mostly simple, occasionally bifurcate. DAGO small, oval, single. Claws striate, not thickened basally. Pulvilli medium, adpressed. Parempodia flap-like, convergent apically. Tricho- bothria 6:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.56. Head w. 1.04, 1. 0.82, vertex 0.42. Antenna total 1. 3.45, I 0.43,0.38, II 1.56, 1.42, III 0.82, 0.79, IV 0.64. Rostrum 10.36, IV 0.46, 0.42. Pronotum a.w. 0.86, p.w. 1.06, 1. 0.60. Wing pad 0.88.Metafemur 1. 1.50, 1.40, w. 0.34, 0.32. Metatibia 1. 2.22, 2.16. Metatarsus I 0.18, 0.17, II 0.42, 0.44. DAGO 0.04.

Material examined: 1 ex., Herts, Rothamsted, 5.VII.48, TRES, (exuvium of Vth instar). 1 ex., London, East Peckham, 6.VII.48, TRES.

Remarks: The nymphs of this species were previously des- cribed by Butler (1923) and illustrated by Kullenberg (1944). - 360 - Heterocordylus genistae Scopoli Figs. 30:8; 59

Blackish-brown. Antennal segments 1,I1,III blackish- brown; segment IV reddish-brown. Head and nota blackish- brown with pale red ecdysial lines. Eyes blackish-brown. Abdomen red, segments transversely marked with red, pale ochreous and blackish-brown stripes and with dorsal and ventral rows of black spots laterally. Posterior two abdominal segments blackish-brown. Antennal segments I and II slightly incrassate. Rostrum reaching nearly to mesothoracic coxae. Eyes touching pronotum. Dorsal body hairs short, curved, simple or longer trifurcate, arising from minute abdominal spots. DAGO oval, single, darkly sclerotised with anterior bar straight. Claws striate, slightly curved apically. Pulvilli medium, rugose, detached from claw apically. Parempodia flap-like, striate, truncate apically. Trichobothria 6:5, mesofemoral trichs long, trichomae present, compact. Bothria recessed.

Measurements: Total body 1. 2.92-3.40 (3.16) n=6. Head w. 0.68-0.96 (0.91) n=6, 1.0.56-0.62.(0.59) n=6, vertex 0.50-0.56 (0.52) n=6. Antenna total 1. 1.92-2.06 (1.99) n=5, I 0.30-0.34 (0.32) n=6, II 0.76-0.86 (0.81) n=6, III 0.44-0.50 (0.46) n=5, IV 0.38-0.40 (0.39) n=5. Rostrum total 1. 1.06-1.18 (1.15) n=6, I 0.30-0.33 (0.32) n=4, II 0.28-0.30 (0.30) n=5, III 0.20-0.25 (0.23) n=4, IV 0.28- 0.30 (0.29) n=4. Pronotum a.w. 0.76-0.88 (0.81)n=6, p.w. 1.05-1.18 (1.09) n=5, 1. 0.48-0.50 (0.48) n=6. Wing pad 0.50-0.74 (0.61) n=6. Metafemur 1.0.88-0.98 (0.92) n=6, w. 0.28-0.33 (0.30) n=5. Metatibia 1. 1.18-1.30 (1.23) n=6. Metatarsus I 0.14-0.15 (0.14) n=6, II 0.34-0.38 (0.36) n=5.

Material examined: 6 ex., Sussex, Horsham Road, 15.VI.25 EAB.

Remarks: Butler did not give the host plant for these speci- mens, but it is likely to be Genista tinctoria. Brown (1924) believed this species to be primarily phytophagous and des- cribed the egg immature stages and life history. Butler (1924) added a description of the early instars. - 361 -

Heterocordylus tibialis (Hahn) Figs. 19:112,113; 31:1; 59

Blackish-brown and red. Antennae blackish-brown. Head and nota blackish-brown with pale ochreous ecdysial lines. Eyes dark red. Abdominal segments red with transverse ochreous and black bands and lateral, dorsal and ventral brown to blackish-brown spots. Posterior two abdominal segments dorsally blackish-brown, colouration extending ventrally to the Xth segment in ð, IXth segment in Coxal margins black. Femora and tibiae brown to blackish-brown. Tarsi black. Antennal segment II sometimes slightly incrass- ate apically in ?. Head and nota rugulose, shiny. Dorsal surface hairs short, curved, simple; longer bifurcate and trifurcate hairs arising from minute, dark, abdominal spots. DAGO single, sclerotised with anterior bar straight. Claws striate. Pulvilli medium, detached apically. Parempodia flap-like, striate, truncate apically. Trichobothria 6:6, trichomae present compact. Bothria recessed.

Measurements: Total body 1. 3.48-3.80 (3.64) n=8. Head w. 0.96-1.10 (1.01) n=8, 1. 0.66-0.76 (0.69) n=8, vertex 0.52- 0.60 (0.56) n=8. Antenna total 1. 2.12-2.40 (2.22) n=8, I 0.30-0.40 (0.36) n=8, II 0.82-1.00 (0.94) n=8, III 0.52- 0.60 (0.56) n=8, IV 0.36-0.40 (0.38) n=8. Rostrum total 1. 1.25-1.30 (1.28) n=5, I 0.30-0.40 (0.35) n=4, II 0.28- 0.32 (0.30) n=4, III 0.24-0.30 (0.26) n=6, IV 0.32-0.34 (0.33) n=7. Pronotum a.w. 0.96-1.04 (1.00) n=8, p.w. 1.18- 1.30 (1.25) n=8, 1. 0.52-0.60 (0.55) n=8. Wing pad 0.54- 0.70 (0.63) n=8. Metafemur 1. 0.92-1.08 (1.01) n=8, w. 0.30- 0.34 (0.32) n=8. Metatibia 1. 1.30-1.48 (1.40) n=8. Meta- tarsus I 0.14-0.16 (0.16) n=8, II 0.39-0.42 (0.40) n=8.

Material examined: 2 ex., Herts, Harpenden, 12.VII.54, Sarothamnus scoparius, TRES. 11 ex., Berks, Silwood Park, 23.VI.77, S. scoparius, GMcG. 10 ex., Sutherland, Bonar Bridge, 5.VII.77, S. scoparius, RGP. 7 ex., Kent, Blean Wood, 10.VII.78, S. scoparius, WRD.

Remarks: Specimens collected at Bonar Bridge were darker than the Silwood Park specimens. This species differs from - 362 - the preceding one in the number of trichobothria, the relative lengths of the mesofemoral trichs and the degree of thickening of the second antennal segment. The occur- rence on broom and the biology of the immature stages have been discussed by Waloff and Southwood (1960). - 363 - Heterotoma planicornis (Pallas) Figs. 7a; 17b-d; 19:114,115; 31:2; 42a,d; Pls 1; 2

Reddish-brown. Antennal segments I, II reddish-brown, III pale, IV with red tinge. Head and nota reddish-brown, eyes red. Abdomen dark red or greenish-red, legs greenish- yellow. Tarsal segment II with apical half infuscate. Anten- nal segments I, II greatly incrassate with dense,black, lanceolate pubescence, III, IV slender with weak pubescence. Rostrum reaching just to metathoracic coxae. Pronotum with lateral margins sub-parallel. Dorsal body hairs black, bifurcate, trifurcate and tapering slightly basally (max.l. 0.14). Femora and ventral body surface with hairs pale, single. Tibiae with pale, weak spines. DAGO small, oval, single with bright red patch anteriad. Claws slender,striate. Pulvilli small, to medium. Parempodia striate, large, flap- like, truncate apically. Trichobothria 7:6, mesofemoral trichs long, trichomae present. Bothria recessed.

Measurements: Total body 1. 3.44-3.76 (3.61). Head w. 0.76-0.84 (0.81), 1. 0.64-0.72 (0.68), vertex 0.38-0.44 (0.41). Antenna total 1. 2.76-2.92 (2.83), I 0.50-0.54 (0.52), II 1.36-1.46 (1.39), III 0.40-0.44 (0.42), IV 0.46- 0.52 (0.49). Rostrum total 1. 1.30-1.46 (1.39), I 0.26- 0.36 (0.33), II 0.32-0.40 (0.36), III 0.30-0.38 (0.33), IV 0.34-0.40 (0.36). Pronotum a.w. 0.66-0.74 (0.71), p.w. 0.82-0.94 (0.86), 1. 0.42-0.48 (0.45). Wing pad 0.56-0.70 • (0.65). Metafemur 1. 1.20-1.32 (1.26), w. 0.30-0.34 (0.32). Metatibia 1. 1.70-1.86 (1.80). Metatarsus I 0.14-0.16 (0.15), II 0.42-0.54 (0.44) . n=10 in all cases.

Material examined: 3 ex., Oxon, Hartswood, 27.VII.48, TRES. 10 ex., London, Hampstead Heath, 25.VI.77, Lamium sp., Crataegus sp., GMcG. 3 ex., London, Putney Heath, 27.VII.77, Quercus sp., GMcG. 8 ex., Bucks, Denham, 29.VII.77, Rubus sp., GMcG.

Remarks: This common speciestas been described by Butler(1923), Van Dinther (1953), Southwood and Scudder (1956b)and Waloff and Southwood (1960) included it in a key to mirid nymphs occurring on Sarothamnus scoparius. The nymphs are easily - 364 - recognisable by the greatly swollen first and second anten- nal segments. The function of this peculiar structure is not known, and although SEM studies have not given any clue as to what this might be, histology has shown that they are probably not at all heavy (there appear to be large intra- cellular spaces). When disturbed, the nymphs either wave their antennae about in slow, circles, or remain motionless with the antennae held at approximately 120° apart. This may have the effect of making the insect appear larger or more frightening to a predator. Swelling of antennal seg- ments occurs in other mirid species, e.g. Atractotomus mali, but never to the same degree. When probing the surface of leaves, antennal segments I and II are held out horizontally, segments III and IV being bent at right angles,with the apex of the IVth segment touching the leaf surface. - 365 -

Blepharidopterus angulatus (Fallen) Figs. la; 17e; 19:116,117; 31:3; 41b; 43c

Light green or greenish-yellow. Antennal segment I with infuscate annulation basally. Head, nota and abdomen light green or greenish-yellow, posterior abdominal segment black. Tibiae with proximal apices black. Apical half of tarsal segment II infuscate. Dorsal body hairs dark, simple, long (max.i. 0.13). Ventral body hairs simple. DAGO small, oval, single with bright orange patch anteriad. Claws slender, striates Pulvilli small to medium. Parempodia large, flap- like, truncate. Trichobothria 5:5, trichomae absent. Bothria slightly tuberculate.

Measurements: Total body 1. 3.52-4.08 (3.87). Head w. 0.70- 0.80 (0.75), 1. 0.60-0.64 (0.61), vertex 0.39-0.44 (0.41). Antenna total 1. 3.61-3.92 (3.71), I 0.56-0.66 (0.60), II 1.22-1.38 (1.28), III 1.18-1.34 (1.24), IV 0.56-0.64 (0.59). Rostrum total 1. 1.22-1.38 (1.31) n=9, IV 0.34-0.36 (0.35) . Pronotum a.w. 0.61-0.70 (0.65), p.w. 0.77-0.88 (0.82), 1. 0.42-0.48 (0.44). Wing pad 0.52-0.72 (0.60). Metafemur 1. 1.42-1.52 (1.47), w. 0.23-0.30 (0.27). Metatibia 1. 2.08- 2.28 (2.18). Metatarsus I 0.14-0.16 (0.15), II 0.36-0.45 (0.41). n=10 except where shown.

Material examined: 20 ex., Oxon, Hambledon, 8.VII.48, Alnus sp., TRES. 15 ex., London, Hampstead Heath, 2.VII.76, Betula sp., GMcG. 18 ex., London, Hampton Court, 11.VII.76, Alnus sp., GMcG. 8 ex., Bucks, Denham, 29.VII.77, Corylus avellana, MMM.

Remarks: The nymphs and adults of this species, which was previously described by Butler (1923), are readily recog- nisable by their black 'knees'. - 366 -

Pachylops bicolor (Douglas and Scott) Figs. 19:118,119; 31:4; 57

Greyish-green. Claws and apex of rostral segment IV infus- cate. Rostral segments III, IV short, squat. Dorsal body hairs stout, simple, black, bifurcate or trifurcate arising from small black spots. Ventral body hairs pale, slender, simple. DAGO single, anterior bar straight, with sinuate membrane in the opening. Claws striate. Pulvilli small. Parempodia large, flap-like. Trichobothria 8:7, trichomae present. Bothria recessed.

Measurements: Total body 1. 2.77-3.04 (2.88). Head w. 0.73- 0.79 (0.76), 1. 0.44-0.54 (0.50), vertex 0.40-0.46 (0.43). •Antenna total 1. 1.98-2.32 (2.12), I 0.26-0.28 (0.27 ), II 0.68-0.84 (0.75), III 0.64-0.82 (0.71), IV 0.37-0.40 (0.39). Rostrum total 1. 0.93-1.02 (0.96), I 0.30-0.34 (0.32), II 0.26-0.30 (0.27), III 0.12-0.14 (0.13), IV 0.22-0.24 (0.23). Pronotum a.w. 0.60-0.70 (0.66), p.w. 0.82-0.86 (0.84), 1. 0.34-0.37 (0.36). Wing pad 0.48-0.62 (0.56). Metafemur 1. 0.84-1.01 (0.91), w. 0.27-0.32 (0.30). Metatibia 1. 1.22- 1.46 (1.29). Metatarsus I 0.11-0.14 (0.12), II 0.30-0.38 (0.35). n=10 in all cases.

Material examined: 5 ex., Herts, Harpenden, 1.IX.54, Sarothamnus scoparius, TRES. 9 ex., Herts, Harpenden, 26.VII.55, TRES. 2 ex., Berks, Silwood Park, 2.VIII.56,TRES.

Remarks: Butler (1923) described the Vth instar nymphs and gave Ulex sp., as the major host plant. The wing pads appear relatively long in this species, their apices darken- ing very considerably prior to the imaginal moult. The interommatidial setae are quite numerous and long. - 367 - Orthotylus flavinervis (Kirschbaum) (S. Orthotylus s . s . ) Figs. 19:124,125; 31:5; 58

Green. Eyes reddish-brown. Tarsi, claws infuscate. Dorsal body hairs mostly short, simple, occasionally longish, bifurcate. Rostrum segment IV with apical setae not highly curled. Tibial spines stout. DAGO single, pale. Claws thick basally, striate. Pulvilli small. Parempodia flap-like, striate. Trichobothria 7:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.32-4.68 (4.56). Head w. 0.88- 0.92 (0.89), 1. 0.68-0.76 (0.72) vertex 0.52-0.56 (0.54) . Antenna total 1. 2.80-3.12 (2.94) n=5, I 0.40-0.44 (0.41), II 1.14-1.28 (1.22), III 0.72-0.82 (0.75) n=5, IV 0.50-0.58 (0.54) n=5. Rostrum total 1. 1.32-1.42 (1.35), I 0.30-0.36 (0.33)n=5, II 0.34-0.38 (0.36) n=5, III 0.29-0.32 (0.31) n=5, IV 0.36 (0.36). Pronotum a.w. 0.80-0.90 (0.85), p.w. 1.10-1.18 (1.13), 1. 0.57-0.60 (0.58). Wing pad 0.46-0.60 (0.51). Metafemur 1. 1.38-1.44 (1.41), w. 0.32-0.36 (0.34). Metatibia 1. 1.94-2.04 (1.99). Metatarsus I 0.16-0.18 (0.17), II 0.44-0.48 (0.46). DAGO 0.086-0.088 (0.087). n=6 except where shown.

Material examined: 6 ex., Herts, Harpenden, 15.VII.54, Acer pseudoplatanus, TRES. 8 ex., Oxon, Ditton Park Station, ll.VT.61, Alnus sp., GEW. - 368 -

Orthotylus ochrotrichus Fieber (S. Orthotylus s.s.) Fig. 58

Light green. Eyes red. Apical portion of tarsi, tip of rostral segment IV infuscate. Dorsal body hairs short, pale, simple, or longish, bifurcate (max.l . 0.10) . Ventral body hairs long, simple. Rostrum segment IV with apical setae very short, highly curled. DAGO small, single, indistinct. Claws striate. Pulvilli medium, rugulose. Parempodia large, flap-like, striate. Trichobothria 7:6, trichomae present, compact. Bothria recessed.

Measurements: Total body 1. 3.20-3.76 (3.42) n=10. Head w. 0.72-0.78 (0.74)n=9, 1. 0.56-0.64 (0.60) n=10, vertex 0.40- 0.44 (0.42) n=10. Antenna total 1. 2.84-3.06 (2.96) n=8, I 0.36-0.40 (0.38) n=10, II 1.10-1.26 (1.18) n=10, III 0.84-0.96 (0.90) n=10, IV 0.50-0.58 (0.53) n=8. Rostrum total 1. 1.33-1.43 (1.38) n=8, IV 0.38-0.40 (0.39) n=9. Pronotum a.w. 0.59-0.71 (0.66) n=10, p.w. 0.76-0.88 (0.82) n=10, 1. 0.38-0.44 (0.40) n=10. Wing pad 0.56-0.70 (0.62) n=10. Metafemur 1. 1.26-1.40 (1.31) n=10, w. 0.24-0.32 (0.28) n=10. Metatibia 1. 1.86-2.14 (1.99) n=10. Metatarsus 10.13-0.14 (0.14) n=9, II 0.39-0.42 (0.40) n=9.

Material examined: 6 ex., Herts, Harpenden, -.VII.54, Urtica dioica , TRES. 4 ex., Herts, Harpenden, 5.VIII.54, TRES.

Remarks: This species is easily confused with Orthotylus prasinus (Southwood & Leston, 1959), and has been previously described by Southwood and Scudder (1956b). - 369 - Orthotylus prasinus (Fallen) (S. Orthotylus s.s.) Figs. 19:130,131; 58

Light green. Dorsal body hairs pale, simple (max.l. 0.11) and bifurcate (max.l. 0.06). Rostral segment IV with setae highly curled. Tibial spines shorter than tibial breadth. DAGO small, oval, single. Claws striate, thick basally. Pulvilli small to medium. Parempodia large, truncate, flap-like. Trichobothria 7:6, trichomae marked, compact. Bothria recessed.

Measurements: Total body 1. 3.52-3.85 (3.66) n=6. Head w. 0.74-0.76 (0.74) n=5, 1. 0.58-0.62 (0.60) n=6, vertex 0.41- 0.46 (0.44) n=5. Antenna total 1. 3.22-3.48 (3.34) n=6, 10.40-0.44 (0.42) n=6, II 1.28-1.36 (1.32) n=6, III 0.93- 1.00 (0.95) n=6, IV 0.61-0.66 (0.64) n=6. Rostrum total 1. 1.40-1.48 (1.43) n=6, I 0.33-0.34 (0.34) n=3, II 0.31-0.34 (0.32) n=3, III 0.36 (0.36) n=3, IV 0.40 (0.40) n=6. Pronotum a.w. 0.68-0.70 (0.69) n=5, pronotum p.w. 0.83- 0.88 (0.85) n=5, 1. 0.38-0.44 (0.41) n=6. Wing pad 0.60- 0.66 (0.63) n=5. Metafemur 1. 1.36-1.50 (1.41) n=6, w. 0.32-0.35 (0.33) n=6. Metatibia 1. 2.06-2.32 (2.19) n=6. Metatarsus I 0.13-0.14 (0.14) n=6, II 0.38-0.40 (0.39) n=6.

Material examined: 18 ex., Westm, Brigsteer,26.VII.54, Corylus avellana , TRES.

Remarks: The apical,prothoracic, tibial spines are used in the key to species. - 370 - Orthotylus ericetorum (Fallen) (S. Litocoris)

Yellowish-green. Eyes red. Tarsi and apex of rostral seg- ment IV infuscate. Dorsal body hairs short or long, black, simple or slightly truncate apically. Antennal segment I with 1 or 2 stout spines. Rostral segment IV with apical setae not very curled. Tibiae with stout spines. DAGO oval, single with orange patch anteriad. Claws thick basally, striate. Pulvilli small. Parempodia flap-like,truncate. Trichobothria 7:6, trichomae marked, compact. Bothria re- cessed.

Measurements: Total body 1. 2.64-2.95 (2.78) n=9. Head w. 0.66-0.70 (0.68) n=9, 1. 0.50-0.56 (0.53) n=9, vertex 0.40-0.44 (0.42) n=9. Antenna total 1. 2.37-2.40 (2.40) n=7, I 0.27-0.30 (0.29) n=9, II 0.86-0.90 (0.87) n=9, III 0.68-0.70 (0.68) n=9, IV 0.52-0.56 (0.54) n=7. Rostrum total 1. 1.18-1.36 (1.28) n=9, IV 0.34-0.38 (0.36) n=8. Pronotum a.w. 0.58-0.66 (0.62) n=8, p.w. 0.70-0.80 (0.73) n=8, 1. 0.32-0.37 (0.34) n=9. Wing pad 0.46-0.54 (0.51) n=9. Metafemur 1. 1.05-1.10 (1.06) n=8, w. 0.28-0.32 (0.30) n=8. Metatibia 1. 1.34-1.49 (1.42) n=8. Metatarsus I 0.12- 0.14 (0.13) n=8, II o.31-0.36 (0.34) n=8.

Material examined: 20 ex., Westm, Foulshaw Moss, 26.VI.54, Calluna vulgaris, TRES. 3 ex., Kent, Perrywood, 26.VII.56, C. vulgaris, TRES. 9 ex., Surrey, Longcross, 20.VII.77, C. vulgaris, GMcG.

Remarks: One of 9 Vth instar nymphs caught at Longcross was parasitised by a braconid sp. The apical spines on the prothoracic tibiae are used in the key to species. - 371 -

Orthotylus adenocarpi (Perris) (S. Neopachylops) Figs. 19:122,123; 58

Green. Head, nota , appendages olive green. Tarsi infuscate. Dorsal body hairs longish, black, scattered, simple. Ventral body hairs pale, simple. Antennal segment I with 3 stout spines. Rostrum segment IV infuscate with apical setae slightly curled. DAGO small, oval, single. Claws striate. Pulvilli small Parempodia large, flap-like, striate, trun- cate. Trichobothria 7:6, trichomae present. Bothria recesssed.

Measurements: Total body 1. 2.80-3.30 (3.02). Head w. 0.72-0.78 (0.75) n=8, 1. 0.49-0.58 (0.52) n=9, vertex 0.40- 0.43 (0.41) n=6. Antenna total 1. 2.05-2.35 (2.18), I 0.29-0.32 (0.30), II 0.82-0.91 (0.88), III 0.52-0.61 (0.58), IV 0.41-0.43 (0.42). Rostrum total 1. 1.20-1.22 (1.21), IV 0.30-0.32 (0.31). Pronotum a.w. 0.60-0.70 (0.66), p.w. 0.80-0.90 (0.85), 1. 0.37-0.44 (0.41). Wing pad 0.48- 0.62 (0.53). Metafemur 1. 0.96-1.04 (1.00), w. 0.30-0.34 (0.32). Metatibia 1. 1.36-1.56 (1.43). Metatarsus I 0.12- 0.14 (0.13), II 0.34-0.39 (0.37). n =10 except where shown.

Material examined: 12 ex, Berks, Silwood Park, 2.VII.56, Sarothamnus scoparius, TRES.

Remarks: The biology and occurrence of the nymphs of this species have been described by Waloff and Southwood (1960). The apical spines on the prothoracic tibiae are used in the key to species. - 372 - (Kirschbaum) (S. Neopachylops) Fig. 19:128,129

Green. Head, nota , appendages olive green. Dorsal body hairs dense, long, dark, bifurcate, trifurcate or coronate (max.l. 0.09) and shorter, dark, simple (max.l. 0.05). Ventral body hairs long, simple (max.1. 0.11). Rostrum segment IV with apical setae straight. DAGO small, oval, single. Claws striate, slightly curved apically. Pulvilli small to medium. Parempodia large, flap-like. Trichobothria 7:6, trichomae marked, compact. Bothria recessed.

Measurements: Total body 1. 2.84-3.20 (3.07) n=4. Head w. 0.74-0.76 (0.75) n=4, 1. 0.56-0.60 (0.57) n=4, vertex 0.43- 0.47 (0.46) n=3. Antenna total 1. 2.32-2.52 (2.43) n=2, I 0.30-0.36 (0.32) n=4, II 0.84-0.92 (0.89) n=4, III 0.78- 0.90 (0.84) n=2, IV 0.39-0.41 (0.40) n=3. Rostrum total 1. 1.06-1.12 (1.08) n=4, IV 0.27-0.28 (0.27) n=4. Pronotum a.w. 0.62-0.70 (0.67) n=4, p.w. 0.78-0.84 (0.81) n=4, 1. 0.38- 0.40 (0.39) n=4. Wing pad 0.45-0.66 (0.57) n=4. Metafemur 1. 1.04-1.10 (1.06) n=4, w. 0.29-0.34 (0.31) n=4. Meta- tibia 1. 1.48-1.64 (1.54) n=4. Metatarsus I 0.13-0.14 (0.14) n=4, II 0.38-0.44 (0.40) n=4.

Material examined: 4 ex., Berks, Silwood Park, 8.VIII.56, Sarothamnus scoparius, TRES.

Remarks: The biology and occurrence of the nymphs of this species on broom hare been described by Waloff and Southwood (1960). One of the 4 nymphs above was parasitised by a braconid sp. The apical spines of the prothoracic tibiae are used in the key to species. - 373 - Orthotylus virescens (Douglas and Scott) (S. Neopachylops) Figs. 19:126,127; 58

Green. Head, nota olive green with broad,white ecdysial lines. Legs green. Tarsi, rostral segment IV slightly in- fuscate. Claws black. Rostrum short,squat, joint between segments III and IV oblique. Dorsal body hairs dense, long, black, simple, bifurcate or occasionally trifurcate, often arising from dark spots.(max.l. 0.11). Ventral body hairs simple. DAGO small, oval, single. Claws striate, thick bas- ally. Pulvilli small. Parempodia large flap-like, striate. Trichobothria 7:6, trichomae marked, compact. Bothria reces- sed.

Measurements: Total body 1. 3.16-3.56 (3.34) n=6. Head w. 0.70-0.76 (0.74), 1. 0.58-0.60 (0.59), vertex 0.30-0.42 (0.40) n=6. Antenna total 1. 2.48-2.85 (2.65) n=6, I 0.30- 0.34 (0.33), II 0.88-1.02 (0.95), III 0.86-1.03 (0.95), IV 0.41-0.46 (0.44) n=6. Rostrum total 1. 0.82-0.86 (0.85) n=5, IV 0.21-0.24 (0.22). Pronotum a.w. 0.64-0.70 (0.67), p.w. 0.82-0.90 (0.87), 1. 0.37-0.42 (0.40). Wing pad 0.50- 0.65 (0.60). Metafemur 1. 1.06 -1.18 (1.11), w. 0.27-0.33 (0.30). Metatibia 1. 1.54-1.80 (1.67). Metatarsus 10.12- 0.14 (0.13), II 0.38-0.44 (0.41). n=7 except where shown.

Material examined: 7 ex., Herts, Harpenden, 3.VII.54, Sarothamnus scoparius, TRES. 15 ex., Midlothian, Edinburgh; 11.VII.78, S. scoparius, GMcG.

Remarks: The biology and occurrence of the nymphs of this species on broom have been described by Waloff and Southwood (1960), and a brief description was given by Butler (1923). This species is the only Orthotylus species on broom to have long dorsal hairs arising from black spots. The articu- lation between the IIIrd and IVth rostral segments is pecu- liar in that it is oblique in relation to the long axis of the rostrum and not transverse as is normally the case. - 374 -

Orthotylus flavosparsus (Sahlberg) (S. Melanotrichus) Figs. 19:120,121; 58

Bright green. Apex of rostral segment IV, apical half of tarsi infuscate. Rostrum short with rostral segment IV squat with joint transverse. Dorsal body hairs fine, black, bifurcate or simple, scattered, arising from minute, dark spots on nota (max.l. 0.08). Rostral segment IV with apical setae straight. DAGO small, oval, single. Claws striate, not very curved. Pulvilli small. Parempodia large, flap- like. Trichobothria 7:6, trichomae present. Bothria recessed..

Measurements: Total body 1. 2.80-3.16 (2.96) n=7. Head w. 0.68-0.76 (0.72) n=8, 1. 0.51-0.60 (0.55) n=8, vertex 0.38- 0.44 (0.41) n=7. Antenna total 1. 0.50-0.62 (0.56) n=7, I 0.28-0.32 (0.30) n=8, II 0.82-0.90 (0.85) n=8, III 0.86- 0.92 (0.88) n=8, IV 0.50-0.54 (0.52) n=7. Rostrum total 1. 0.88-1.00 (0.93) n=7, IV 0.21-0.24 (0.22) n=8. Pronotum a.w. 0.58-0.68 (0.64) n=7, p.w. 0.79-0.88 (0.82) n=8, 1. 0.34-0.40 (0.37) n=7. Wing pad 0.50-0.60 (0.55) n=8. Metafemur 1. 0.98-1.04 (1.01) n=8, w. 0.30-0.37 (0.33) n=8. Metatibial 1. 1.40-1.49 (1.44) n=8. Metatarsus I 0.12 -0.13 (0.12) n=7, II 0.39-0.42 (0.41) n=7.

Material examined: 10 ex., (no data), TRES.

Remarks: Although the host plant was not recorded for the above specimens, this species, described by Reuter (in Butler, 1923), is well known to be abundant on members of the Chenopodiaceae. The apical spines of the prothoracic tibiae are used in the key to species. - 375-

Orthotylus moncreaffi (Douglas and Scott) (S. Melanotrichus)

Green. Rostrum segment IV with apex infuscate. Dorsal body hairs dark, simple (max.i. 0.067). Ventral body hairs slender, simple. Rostrum segment IV with apical setae slightly curled. DAGO small, oval, single. Claws striate. Pulvilli small. Parempodia large, flap-like. Trichobothria 7:6, trichomae present, compact. Bothria recessed.

Measurements: Total body 1. 2.60-2.80 (2.67). Head w. 0.72- 0.80 (0.75), 1. 0.52-0.57 (0.55), vertex 0.40-0.50 (0.44) . Antenna total 1. 2.06-2.16 (2.10), I 0.25-0.30 (0.27), II 0.70-0.78 (0.75), III 0.64-0.69 (0.66), IV 0.40-0.45 (0.43) . Rostrum total 1. 1.06-1.22 (1.14), IV 0.27-0.30 (0.39). Pronotum a.w. 0.58-0.67 (0.62), p.w. 0.72-0.82 (0.77), 1. 0.30-0.35 (0.33). Wing pad 0.40-0.44 (0.43). Metafemur 1. 0.89-0.96 (0.93), w. 0.28-0.37 (0.32). Metatibia 1. 1.18- 1.32 (1.25). Metatarsus I 0.12-0.13 (0.13), II 0.36-0.42 (0.40). n=9 in all cases.

Material examined: 9 ex., Hants, Pennington, 16.VII.51, TRES.

Remarks: The host plant for the above specimens was not recorded. Southwood and Leston (1959) state ' Although sometimes taken with Orthotylus rubidus and recorded from the same counties, this species usually occurs on larger chenopods, principally sea purslane, often around the high water mark, spring tides The larvae of the two species can be distingiithed by their bolour, those of O. rubidus being pink-red and those of 0. moncreaffi green.' The apical spurs on the prothoracic tibiae are used in the key to species. - 376 - Pseudoloxops coccineus (Meyer-DUr) Figs. 19:132,133; 31:6

Green. Head with lateral crimson stripes from base of antennae, through eyes, continuing down lateral margins of nota . Eyes red. Legs yellow, femora with orange tinge apically, simple. Dorsal body hairs long, black (max.1. 0.22). Ventral body hairs shorter, less stout, simple. Antennal segment I and apex of metafemur with stout hairs. Tibiae with long, spinous, simple hairs, those of meta- tibiae up to 0.17mm. DAGO single, pale. Claws striate, thick basally. Pulvilli indistinct. Parempodia large, flap- like, truncate. Trichobothria 7:6, trichomae very weak. Bothria slightly recessed.

Measurements: Total body 1. 3.36-3.60 (3.47). Head w. 0.72- 0.76 (0.75), 1. 0.56-0.58 (0.57), vertex 0.43-0.48 (0.45). Antenna total 1. 2.62-2.74 (2.69), 10.41-0.42 (0.42), II 1.17-1.22 (1.20), III 0.58-0.60 (0.59), IV 0.46-0.50 (0.48). Rostrum total 1. 1.04-1.12 (1.09), IV 0.29-0.30 (0.30). Pronotum a.w. 0.70-0.74 (0.71), p.w. 0.90-0.96 (0.93), 1. 0.40-0.42 (0.41) . Wing pad 0.54-0.56 (0.55) . Metafemur 1. 1.10-1.20 (1.15), w. 0.35-0.36 (0.36). Metatibia 1. 1.56- 1.60 (1.58). Metatarsus I 0.15-0.16 (0.15), II 0.36-0.38 (0.37). n=3 in all cases.

Material examined: 3 ex., Berks, Ascot, 3.VIII.61, Fraxinus excelsior,TRES.

Remarks: This species, which was described by Butler (1923), is parisitised by a species of braconid. In one slide mount the mouth parts the larval parasite could be seen, but not well enough to enable•a specific or generic identi- fication. - 377 - Cyrtorhinus caricis (Fallen) Figs. 19:134,135; 31:7

Green or orangish-yellow. Eyes red, bordered white laterally. Posterior margins of nota outlined in black. Rostrum reaching mesocoxae. Dorsal body hairs simple, black, occasionally bifurcate (max.l., abdomen 0.07; head and nota 0.09). DAGO single with anterior bar curved. Claws slender, striate, slightly curved. Pulvilli very small. Parempodia flap-like. Trichobothria 7:6, trichs long, tri- chomae strong, compact. Bothria well-recessed.

Measurements: Total body 1. 2.80-3.18 (2.94) n=4. Head w. 0.68-0.77 (0.73) n=4. 1. 0.46-0.50 (0.47) n=4, vertex 0.30- 0.39 (0.36) n=4. Antenna total 1. 2.33, 2.34 (2.34) n=2, I 0.32-0.38 (0.35) n=4, II 0.76-0.78 (0.77) n=3, III 0.70, 0.70 (0.70) n=2, IV 0.50-0.53 (0.51) n=2. Rostrum total 1. 1.02-1.06 (1.04) n=4, IV 0.24-0.26 (0.25) n=3. Pronotum a.w. 0.54-0.58 (0.56) n=3, p.w. 0.74-0.79 (0.76) n=3, 1. 0.36-0.39 (0.37) n=4. Wing pad 0.42-0.46 (0.44) n=3. Metafemur 1. 1.00-1.08 (1.03) n=4, w. 0.20-0.27 (0.22) n=4. Metatibia 1. 1.26-1.35 (1.30) n=4. Metatarsus I 0.13-0.14 (0.14) n=4, II 0.40-0.43 (0.41) n=4. DAGO 0.064.

Material examined: 1 ex., Essex, Epping Forest, 13.VII.12, EAB. 2 ex., Berks, Silwood Park, 27.VII.60, TRES. 1 ex., Dumfs, Bodesbeck Farm, 10.VII.78, Juncus sp., GMcG & MMM.

Remarks: This species is to be found at the base of Carex and Juncus spp., and is known to be predacious on the eggs of delphacids (Southwood & Leston, 1959). Butler (1923) gave a brief description of the nymphs, which are orange in early instars (Southwood & Leston,1959). Rothschild (1963) studied the biology of this species in Juncus areas and reported it feeding on the plant stem and most stages of the delphacid Conomelus anceps. A drawing of the Vth instar nymph has been given (Rothschild,1964). - 378 - Neomecomma bilineatus (Fallen) Figs. 19:136, 137; 31:8

Green. Antennae fuscous. Head with central triangular or diamond shaped fuscous mark between eyes. Posterior margin of pronotum, internal margins and apices of meso- and meta- nota outlined in black, fuscous. Posterior abdominal seg- ment, tarsi, claws and rostral segment IV fuscous. Tibiae yellowish-green. Dorsal body hairs dark, simple (max.l., head 0.10; elsewhere 0.08). Ventral body hairs slender, simple. DAGO oval, single with crenate internal membrane. Claws stout, short, thick basally. Pulvilli small. Parem- podia large, flap-like, striate. Ttichobothria 7:6, trichomae present, weak. Bothria recessed.

Measurements: Total body 1. 3.52-3.78 (3.62). Head w. 0.76- 0.80 (0.77), 1. 0.58-0.66 (0.60), vertex 0.34-0.42 (0.38) n=9. Antenna total 1. 2.68-2.96 (2.84), 10.40-0.43 (0.42), II 0.98-1.10 (1.05), III 0.78-0.90 (0.85), IV 0.47-0.56_ (0.52). Pronotum a.w. 0.63-0.68 (0.66) n=9, p.w. 0.86- 0.92 (0.89) n=9, 1. 0.41-0.44 (0.43). Wing pad 0.62-0.75 (0.69). Metafemur 1. 1.19-1.26 (1.22), w. 0.24-0.28 (0.26). Metatibia 1. 1.70-1.80 (1.78). Metatarsus 10.14-0.16 (0.14), II 0.40-0.42 (0.41) n=9. DAGO 0.078. n=10 except where shown.

Material examined: 18 ex., Herts, Brickett Wood, 15.VII.54, TRES.

Remarks: The host plant is not mentioned for the above specimens, but Southwood and Leston (1959) gave Populus tremula and occasionally P. canescens as the usual host plants. Butler (1923) gave a brief description of the Vth instar nymphs. - 379 -

Mecomuua ambulans (Fallen) (S. Mecouuua s . s . ) Figs. 19:138,139; 32:1

Reddish-green. Antennal segments I, II red, basal half of III white, apical half of III, entire IV lightly infuscate. Head and nota red, slightly shiny. Wing pads becoming infuscate apically, slightly shiny. Femora, tibiae pale reddish-green. Head broad. Eyes large, just touching anterior pronotal margin. Dorsal body hairschrk, short, simple. DAGO single, sclerotised, constricted medially with anterior bar strong. Claws slender, striate. Pulvilli small, Parempodia flap-like. Trichobothria 8:5, trichomae strong, compact. Bothria well-recessed.

Measurements: Total body 1. 3.00-3.20 (3.13).Head w. 0.74- 0.80 (0.77), 1. 0.50-0.54 (0.51), vertex 0.38-0.40 (0.39). Antenna total 1. 2.34-2.87 (2.55), I 0.36-0.42 (0.38), II 0.76-0.94 (0.83), III 0.78-1.00 (0.87), IV 0.40-0.51 (0.46). Rostrum total 1. 1.20, 1.34 (1.27) n=2, IV 0.28-0.34 (0.31). Pronotum a.w. 0.57-0.64 (0.59), p.w. 0.76-0.84 (0.79), 1. 0.36-0.45 (0.40). Wing pad ? 0.44 n=1, 60.74 n=1. Meta- femur 1. 1.06-1.28 (1,16), w. 0.21-0.24 (0.23). Metatibia 1. 1.56-1.88 (1.66) . Metatarsus I 0.14, 0.15 (0.15) n=2, II 0.41,0.48 (0.45) n=2. DAGO 0.080. n=3 except where shown.

Material examined: 1 ex., Herts, Rothamsted, 14.VII.54, TRES. 3 ex., Bucks, Burnham Beeches, 5.VII.58, GEW. 2 ex., Perths, Kinloch Rannoch, 26.VI.78, GMcG.

Remarks: The nymphs of this species have been briefly des- cribed by Butler (1923). Southwood and Leston (1959) gave Juncus and Carex spp. as the food plants of this species. The specimens from Kinloch Rannoch were collected from a damp meadow, with vegetation consisting of grass species, bracken, Ranunculus sp. and Veronica chamaedrys. The body of the ? nymph is broader and more squat, the wing pads extending only just to the DAGO, whereas on the 6 they ex- tend past the DAGO. Antennal segments I and II are slightly incrassate in the d, and more strongly incrassate at the apex of segment II in the ?. There is no trace of the black colouration present in the adults. - 380 -

Myrmecoris gracilis (Sahlberg) Figs. 32:3; 60

Reddish-brown. Antennal segment I, II reddish-brown, seg- ment III with basal one quarter white, apical three- quarters and entire segment IV reddish-brown. Ant-like. Antennal segment II slightly incrassate apically. Head long, eyes large, touching pronotum. Rostrum reaching meso-thoracic coxae. Pronotum square with posterior angles very rounded, with strong constriction posteriad. Meso- and meta-thoracic nota narrow posteriad. Abdominal segments I, II narrow, rest of abdomen becoming full and rounded. DAGO small, single, margins strongly sclerotised. Claws straight. Pulvilli indistinct. Parempodia flap-like. Trichobothria 8:6, trichs long, trichomae compact. Bothria recessed.

Measurements: Total body 1. 4.52. Head w. 0.94, 1. 0.90, vertex 0.40. Antenna total 1. 4.16, 10.30, II 1.48, III 1.28, IV 1.10. Rostrum segment IV 0.38. Pronotum a.w. 0.57, 1. 0.60. Metafemur 1. 1.60. Metatibia 1. 2.16. n=1 in all cases.

Material examined: 1 IVth ex., Surrey, Chobham Common, -eVI.59, GEW.

Remarks: The ant-like form of this predacious species is readily identifiable. It is found in areas of Pinus-Calluna heathland colonised by sheep's fescue and wavy hair grass, and feeds on grass aphids, their , the eggs of Notostira and other small animals (Southwood & Leston,1959). The wing pads of the IVth instar nymph examined were very mall. - 381 - Pithanus maerkeli (Herrich-Schgffer) Figs. 19:142,143; 32:4; 60

Dark reddish-brown or brownish-black. Antennal segment I with basal half reddish-brown, apical half white, II, III, IV light reddish-brown. Head, pronotum dark reddish-brown with bright red ecdysial line. Wing pads pale ochreous apically. Ventral, dorsal abdominal surfaces reddish-brown with lateral margins pale ochreous, except dorsally where broken by transverse, thin, red stripe extending from DAGO. Head short. Pronotum swollen medially, slight constriction anteriad amd posteriad. Wing pad short, blunt. Dorsal body hairs short, simple, occasionally long, blunt on posterior abdominal segments. DAGO double, anterior bar sinuate, sclerotised. Claws slender, nearly straight, striate. Pul- villi indistinct. Parempodia flap-like, striate. Trichobothria 8:6, ventral metathoracic femoral trichs very long, trichomae compact. Bothria recessed.

Measurements: Total body 1. 3.96-4.22 (4.13) n=3. Head w. 0.90-1.00 (0.96), 1. 0.70-0.76 (0.73), vertex 0.44-0.54 (0.47). Antenna total 1. 3.46, 3.76 n=2, I 0.38-0.40 (0.40), II 1.04-1.32 (1.24), III 1.24,1.30 n=2, IV 0.80, 0.82 n=2. Rostrum total 1. 1.62-1.70 (1.66) n=4, IV 0.40-0.44 (0.42). Pronotum a.w. 0.78-0.95 (0.84), p.w. 0.84-0.95 (0.87), 1. 0.50-0.52 (0.51). Wing pad 0.08-0.12 (0.10) n=3. Meta- femur 1. 1.32-1.50 (1.43), w. 0.23-0.28 (0.26). Metatibia 1. 1.64-1.88 (1.80). Metatarsus I 0.32-0.36 (0.33), II 0.36- 0.38 (0.37). n=5 except where shown.

Material examined: 2 ex., Hants, Byfleet, 19.VI.15, EAB. 2 ex., Sussex, Horsham Road, 19.VI.25, EAB. 1 ex., Berks, Silwood Park, 5. VI.56, TRES. 1 ex., Perths, Rannoch School, 24.VI.78, GMcG. 3 ex., Perths, Kinnloch Rannoch, 25.VI.78 GMcG. 1 ex., Dumfs, Bodesbeck.Farm, 10.VII.78, GMcG.

Remarks: This partly predacious species is found among grasses in damp meadows where rushes are growing (Southwood & Leston, 1959). - 382 - Lygus maritimus Wagner Figs. 19:144,145; 32:5

Yellowish-green. Eyes tinged red with fuscous mark post- eriad. Pronotum with two blackish-brown spots equidistant from body midline, propleura with one blackish-brown spot laterally. Mesothoracic notum with one black spot on either side of body midline, furtherapart than pronotal spots. Apices of wing pads brownish-black. Distal margins of femora black. Apical half of rostral segment IV, apical one third of tarsal segment II and claws entirely infuscate. Dorsal body surface, antennae, legs with short, sparse, spinous, blackish-brown hairs. Tibiae with strong dark spines, shorter than tibial diameter. DAGO double, anterior bar sinuate, sclerotised, with pigmented patch anteriad and posteriad. Claws slender, slightly curved, striate. Pulvilli medium. Parempodia flap-like. Trichobothria 7:5, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 3.84-4.32 (3.98) . Head w. 1.00- 1.05 (1.03), 1. 0.78-0.86 (0.80), vertex 0.47-0.54 (0.49). Antenna total 1. 2.86-3.02 (2.94), I 0.40-0.42 (0.41), II 1.14-1.22 (1.18), III 0.72-0.77 (0.75), IV 0.56-0.64 (0.62). Rostrum total 1. 1.82-2.04 (1.92), IV 0.49-0.54 (0.52). Pronotum a.w. 0.90-1.04 (0.96), p.w.1.30-1.40 (1.35), 1. 0.64-0.74 (0.68). Wing pad 0.60-0.80 (0.68). Metafemur 1. 1.30-1.40 (1.34), w. 0.33-0.38 (0.35). Metatibia 1. 1.82- 1.94 (1.88). Metatarsus I 0.18-0.21 (0.19), II 0.54-0.58 (0.56). DAGO 0.09 (0.09) n=3. n=5 except where shown.

Material examined: 5 ex,, Hants, Milford on Sea, 11.VII.55, maritima, TRES. 1 ex., Devon, Wembury, 4.VI.60, Beta maritima, TRES.

Remarks: The arrangement of dark spots on the nota is common to the genus Lygus. - 383 -

Lygus rugulipennis Poppius Figs. 19:146,147; 33:1; 74

Green with irregular light brown markings. Eyes with fuscous marks posteriad, almost meeting in body midline. Pronotum with one medial black spot on either side of midline and light brown irregular markings. Propleura with one black spot adjacent to pronotal margins. Mesonotum with one black spot on either side of body midline. Wing pads irregularly marked with brown. Abdominal segments with light brown transverse markings. Legs green with brown banding on femora and tibial bases. Body broad. Dorsal body hairs stout, black, spinous, simple (max.l. 0.07). Ventral body hairs long, pale, simple (max.1.0.086). Tibiae with stout short spines. DAGO large,double with margins heavily sclerotised, anterior bar sinuate, with black patch anteriad and posteriad. Claws thick basally, partially striate. Pulvilli medium. Parem- podia flap-like. Trichobothria 7:6, trichomae weak, compact. Bothria slightly recessed.

Measurements: Total body 1. 4.04-5.40 (4.78). Head w. 0.98- 1.06 (1.03), 1. 0.77-0.96 (0.84), vertex 0.50-0.58 (0.55), Antenna total 1. 2.96-3.48 (3.25), 10.44-0.48 (0.45), II 1.20-1.40 (1.30), III 0.76-0.94 (0.78), IV 0.68-0.76 (0.73). Rostrum total 1 1.86-2.18 (2.06), 10.46-0.60 (0.55), II 0.46-0.60 (0.53), III 0.38-0.44 (0.41), IV 0.54-0.60 (0.58). , Pronotum a.w. 0.94-1.04 (0.97), p.w. 1.32-1.54 (1.44), 1. 0.64-0.78 (0.73) . Wing pad 0.62-0.80 (0.76) . Metafemur 1. 1.36-1.58 (1.48), w. 0.38-0.46 (0.40). Metatibia 1. 1.96- 2.20 (2.05). Metatarsus I 0.20 •(0.20), II 0.56-0.60 (0.58). DAGO 0.11 (0.11) n=5. n=10 except where shown.

Material examined: 4 ex., Herts, Wheathamsted, 28,VIII.48, Chemopodium sp., TRES. 1 ex., Isle; of Scilly, Bryher, -.VIII.65, GEW. 11 ex., Dmfs, Bodesbeck Farm, 2.IX.76, Avena sativa, GMcG.

Remarks: The life cycle and nomenclature was discussed by Southwood (1956c),and the biology and life history of this species on oats in Scotland has been studied by Stewart (1969). - 384 -

Figure 74

Lygus rugulipennis (Vth instar nymph) - 385 -

This species is an important pest of crops Tn the palaearc- tic region and Southwood and Leston (1959) listed the crops involved. The notal spots present are larger than in L. maritimus and the irregular brown patterning on the nota is more extensive and marked. - 386 -

Liocoris tripustulatus (Fabricius) Figs. 19:148,149; 33:2; 42g; 75

Light green, mottled brown. Antennal segment I with mottled reddish-brown markings, II with broad reddish-brown band basally and apically, III similarly with two reddish-brown bands, IV more or less light reddish-brown. Head, nota mottled brown. Mesonotum with lateral anterior margins infuscate, metanotum with medial posterior margins and api - ces with fuscous markings. Femoral light green with one or two reddish-brown mottled transverse bands distally. Tibiae with two reddish-brown bands proximally, apices infuscate. Apical half of tarsal segment II, claws entirely infuscate. Dorsal body hairs black, simple(max.l. 0.06), stout, bifur- cate, trifurcate or serrate on posterior abdominal segments, Ventral body hairs pale, simple. DAGO double, anterior bar sinuate, pigmented and occasionally with posterior red patch. Claws striate, curved. Pulvilli medium. Parempodia flap- like, truncate. Trichob.othria 9:6, trichomae weak. Bothria redessed.

Measurements: Total body 1. 3.68-4.12 (4.02). Head w. 0.82- 0.90 (0.87), 1. 0.66-0.76 (0.73), vertex 0.42-0.50 (0.46). Antenna total 1. 2.22-2.98 (2.82), 10.42-0.48 (0.45), II 0.96-1.08 (1.02), III 0.70-0.74 (0.71), IV 0.68-0.74(0.70). Rostrum total 1. 1.62-1.80 (1.72), 10.44-0.50 (0.48), II 0.36-0.49 (0.42), III 0.30-0.36 (0.32), IV 0.46-0.50 (0.49). Pronotum a.w. 0.72-0.68 (0.79), p.w. 1.02-1.18 (1.13), 1. 0.56 -0.68 (0.60) . Wing pad 0.44-0.66 (0.58). Metafemur 1. 1.28-1.37 (1.32), w. 0.34-0.38 (0.36). Metatibia 1. 1.82- 1.90 (1.86). Metatarsus I 0.19-0.22 (0.20), II 0.50-0.54 (0.54). DAGO 0.09 (0.09) n=5. n=10 except where shown.

Material examined: 12 ex., Herts, Rothamsted, 11-17.VIII.54, Urtica dioica, TRES. 3 ex., London, Hampstead Heath, 25.VI.76 U. dioica, GMcG.

Remarks: This species was previously described by Butler (1923) and by S uthwood.and Scudder (1956b). The latter work discussed parasitism by a euphorine braconid. - 387 -

Figure 75

Liocoris tripustulatus (Vth instar nymph) - 388 -

Orthops viscicola (futon) Figs. 19:152-153; 33:6

Green. Wing pad apices red. Femora tinged red. Rostral tip and claws black. Dorsal body hairs short, simple, black, curved, occasionally long, stout, bifurcate or trifurcate. Cuticular sculpturing as dense, minute spinules occurring in transverse - pattern on antennal segment I. DAGO pale, single or double, anterior bar curved or slightly sinuate. Claws striate, thick basally. Pulvilli small. Parempodia flap-like, striate. Trichobothria 8:6, trichomae absent. Bothria flush.

Measurements: Total body 1. 3.20-3.72 (3.42). Head w. 0.78- 0.90 (0.83), 1. 0.52-0.66 (0.59), vertex 0.40-0.43 (0.42) n=3. Antenna total 1. 2.38-2.51 (2.43) n=6, 10.32-0.34 (0.34), II 0.99-1.10 (1.05), III 0.58-0.63 (0.60), IV 0.42- 0.48 (0.44) n=6. Rostrum total 1. 1.10-1.26 (1.19), IV 0.28-0.30 (0.29). Pronotum a.w. 0.70-0.78 (0.73), p.w. 1.00- 1.20 (1.07), 1. 0.46-0.54 (0.50). Wing pad 0.48-0.62 (0.57). Metafemur 1. 1.00-1.16 (1.05) n=5, w. 0.28-0.36 (0.32) n=5. Metatibia 1. 1.48-1.66 (1.52). Metatarsus I 0.14-0.16 (0.15) n=5, II 0.39-0.42 (0.40) n=5. DAGO 0.04 (0.04) n=3. n=7 except where shown.

Material examined: 15 ex., Berks, Silwood Park, 3.VI.77, Viscum album, GMcG.

Remarks: Nymphs were collected from a large clump of mistle- toe which was cut from the top of a lime tree. The clump was placed under a plastic sheet to raise the temperature, retain moisture and speed up nymphal development from IIIrd and IVth instars. In 4-7 days, before the mistletoe had completely wilted, numerous Vth instar nymphs and 4 adults were found. The bright red colour of the femora described by Butler(1923) was not marked. The red wing pad apices became heavily infuscate prior to the last moult. - 389 -

Orthops campestris (L.) Figs. 33:7

Green. Antennae light brown, segment IV red. Head, nota green with light fuscous markings. Eye facets red. Meso- nottm with central portion green, wing pads pale ochreous. Pleura with red tinges. Abdomen green with medial red stripe, sometimes faint, and two lateral brownish-red stripes. Claws, apical portion of tarsal segment II, apex of rostral seg- ment IV black. Dorsal body hairs black, simple, occasionally bifurcate on posterior abdominal segments (max.1. 0.05). Ventral body hairs black, slender, simple (max.l. 0.07). Tibiae with spines short, weakly striate. Interonrmatidial sculpturing as weak spinules, setae short, simple. DAGO double, black, anterior bar sclerotised, sinuate. Claws faintly striate. Pulvilli small. Parempodia flap-like, trun- cate. Trichobothria 7:6, trichomae weak. Bothria recessed or slightly recessed.

Measurements: Total body 1. 3.20-3.44 (3.29) n=6. Head w. 0.70-0.83 (0.78) n=5, 1. 0.54-0.62 (0.58) n=4, vertex 0.32- 0.40 (0.38) n=5. Antenna total 1. 1.88-2.00 (1.95) n=5, I 0.25-0.32 (0.30) n=6, II 0.70-0.80 (0.74) n=6, III 0.46- 0.48 (0.47) n=5, IV 0.40-0.44 (0.42) n=5. Rostrum segment IV 0.28-0.32 (0.30) n=4. Pronotum a.w. 0.66-0.78 (0.72) n=5, p.w. 0.92-1.10 (1.02) n=5, 1. 0.50-0.54 (0.52) n=4. Wing pad 0.54-0.66 (0.59) n=6. Metafemur 1. 0.84-0.99 (0.91) n=6, w. 0.18-0.26 (0.22) n=6. Metatibia 1. 1.12- 1.28 (1.20) n=6. Metatarsus 1 0.14-0.15 (0.14) n=6, II 0.36- 0.39 (0.38) n=6. DAGO 0.07 (0.07) n=4.

Material examined: 2 ex., Kent, Orpington, 13.VIII.78, , DOD. 4 ex., London, Kew Gardens, 24.VIII.78, H. gwnniferam, DOD.

Remarks: Some nymphs have little red colouration, others have strongly tinged pleura, pronotum, abdomen and femora. In some specimens the abdominal margins have pale red patches on each segment, in addition to the longitudinal stripe. Butler (1923) described the nymphs and gave Pastinaca sativa a's the most common host plant. - 390 - Lygocorts pabulinus (L.) (S. Lygocoris s.s.) Figs. 19:156-157; 34:1

Green, occasionally with yellow tinge. Eyes red. Rostral apex and tarsi infuscate. Dorsal body hairs black, simple (max.l.0.09). Ventral body hairs, pale, simple. DAGO double, weakly sclerotised, anterior bar sinuate. Claws striate. Pulvilli small. Parempodia flap-like, striate. Trichobothria 8:7, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 4.20-4.84 (4.41). Head w. 0.82- 0.94 (0.88), 1. 0.72-0.80 (0.77), vertex 0.38-0.52 (0.45) n=9. Antenna total 1. 3.66-4.30 (4.11) n=9, I 0.48-0.58 (0.54), II 1.38-1.60 (1.50), III 1.06-1.27 (1.17) n=9, IV 0.84-1.00 (0.94) n=9. Rostrum total 1. 1.62-1.88 (1.77). IV 0.44-0.54 (0.51). Pronotum a.w. 0.76-0.90 (0.85), p.w. 1.04-1.28 (1.17), 1. 0.50-0.66 (0.60). Wing pad 0.60-0.84 (0.70). Metafemur 1. 1.46-1.82 (1.66), w. 0.30-0.40 (0.35). Metatibia 1. 2.24-2.68 (2.50). Metatarsus I 0.17-0.20 (0.18), II 0.47-0.60 (0.54). DAGO 0.07 (0.07). n=10 except where shown.

Material examined: 3 ex., Herts, Rothamsted, 5.VIII.54, Urtica dioica, TRES. 20 ex., Herts, Harpenden, 28.VI.55, U. dioica, TRES. 15 ex., Yorks, Malham Tarn, 21.VI.76, U. dioica, GMcG. 16 ex., Berks, Silwood Park, 19.VI.78, Ribes nigrum, RH.

Remarks: Butler (1923) and Petherbridge and Thorpe (1928) described the biology and life history of the nymphs of this species, recorded as a serious pest of and hops (Massee, 1954). Southwood and Scudder (1956b) published drawings and measurements of nymphal stages and included this species in a key to mirid nymphs found on U. dioica. - 391 - Lygocoris contaminatus (Fallen) (S. ) Figs. 19:158,159; 34:2

Bright green. Rostral apex infuscate. Wing pad apices, terminal abdominal segment infuscate. Femora with two, broad faint transverse fuscous bands apically. Tarsi infuscate. Dorsal body hairs dark, long, curved, striate, simple (max. 1. 0.06) and dark,straight, simple (max.l. 0.08). Ventral body hairs pale, long, slender, simple (max.l. 0.09). Meta- thoracic tibial spines long, arising from dark spots. Cuti- cular sculpturing in form of strong spinules. DAGO double, broad, anterior bar sinuate,sclerotised. Claws thick basally, striate. Pulvilli medium, rugulose. Parempodia flap-like, truncate. Trichobothria 8:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.08-4.76 (4.51). Head w. 0.90- 1.04 (0.99), 1. 0.72-0.86 (0.79), vertex 0.48-0.55 (0.50). Antenna total 1. 3.60-4.08 (3.86), I. 0.50-0.56 (0.53), II 1.40-1.64 (1.53), III 0.96-1.06 (1.01), IV 0.74-0.84 (0.78). Rostrum total 1. 1.70-1.86 (1.79), IV 0.50-0.56 (0.52). Pronotum a.w. 0.84-0.94 (0.89), p.w. 1.10-1.30 (1.21), 1. 0.54-0.64 (0.59). Wing pad 0.42-0.84 (0.59). Metafemur 1. 1.43-1.70 (1.57), w. 0.32-0.40 (0.36) . Metatibia 1. 2.24- 2.56 (2.39). Metatarsus I 0.19-0.21 (0.20), II 0.54-0.58 (0.56). DAGO 0.09 (0.09) n=5. n=10 except where shown.

Material examined: 7 ex., Lancs, Rusland Moss, 23.VI.54, Betula sp., TRES.

Remarks: This species,described by Butler (1923),can be distinguished from L. pabulinus in that the tibial spines arise from dark spots. - 392 -

C3mptozygum pinastri (Fallen) Figs. 19:162,163; 34:8; 76

Light brown with red markings. Antennae light brown, segment I with 3 or 4 bright red patches. Head light brown with reddish-black, longitudinal, medial stripe and with red markings at antennal bases, internal margins of eyes, post- erior margin and on ventral surface. Eyes reddish-black. Rostral segment I with dark red longitudinal, lateral stripe, segment IV with apical half infuscate. Pronotum with calli dark red to black. Wing pads with red markings, apices infuscate, posterior margins black. Abdomen light brown with thin, transverse, dark reddish-black banding, ventral surface not banded, dorsal red colouration not extend- ing ventrally past connexivum. Coxae mottled red. Femora with red spots apically and red patches or flashes basally. Tibiae with longitudinal red patches, sometimes as interrup- ted transverse bands. Tibial apices, apical half of tarsal segment II infuscate. Dorsal body hairs short, dark, curved, simple (max.l. 0.04) and long, dark, bifurcate (max.1. 0.06). Ventral body hairs long, pale, simple (max.l. 0.08). Cuticu- lar sculpturing spinous, black. DAGO double,dark, anterior bar sinuate and with red patch anteriad and posteriad. Claws slightly curved, striate. Pulvilli medium, adpressed. Parempodia flap-like, truncate. Trichobothria 7:6, trichomae very weak. Bothria slightly recessed.

Measurements: Total body 1. 3.00-3.60 (3.33) n=5. Head w. 0.96-1.04 (1.00) n=4, 1. 0.68-0.88 (0.76) n=5, vertex 0.54- 0.58 (0.55) n=4. Antenna total 1. 2.41-2.54 (2.48) n=4, I 0.33-0.34 (0.34) n=6, II 1.04-1.08 (1.06) n=6, III 0.50-0.54 (0.52) n=5, IV 0.53-0.60 (0.56) n=4. Rostrum total 1. 1.56- 1.64 (1.59) n=6, IV 0.40-0.42.(0.41) n=6. Pronotum a.w. 0.84-0.86 (0.85) n=3, p.w. 1.10-1.16 (1.14) n=3, 1. 0.52- 0.56 (0.54) n=5. Wing pad 0.44-0.62 (0.53) n=6. Metafemur 1. 1.02-1.10 (1.07) n=6, w. 0.28-0.30 (0.29) n=6. Meta- tibia 1. 1.42-1.50 (1.47) n=5. Metatarsus I 0.17-0.1. (0.18) n=5, II 0.47-0.50 (0.48) n=4. DAGO 0.06 (0.06) n=3.

Material examined: 7 ex., Surrey, Oxshott, -.VI.55, Pinus sp., TRES. 1 ex., Surrey, Virginia Water, 7.VII.62, Pinus sp., - 3 93 -

Figure 76

Camptozygum pinastri (Vth instar nymph) - 394 -

GEW. 2 ex., Berks, Silwood Park, 3.VII.77, Pinus sylvestris, GMcG .

Remarks: The unique colour and patterning of this nymph, together with its host plant, make it readily identifiable. The degree of colouration varies and continental specimens may be much more black (Butler, 1923). Kullenberg (1944) gave a colour sketch of a Vth instar nymph. - 395 -

Polymerus palustris (Reuter) (S. Poeciloscytus) Figs. 19:166,167; 35:3

Green, shiny, tinged with red. Meso- and meta-nota poster- ior margins fuscous. Tarsi and rostral apex fuscous. Claws black. Dorsal body hairs dense, black, stout, simple, arising from minute black spots (max.l. 0.13). Ventral body hairs long, pale, simple (max.l. 0.14). Tibiae with stout, striate, short, black spines. DAGO double, anterior bar slightly sinuate. Claws faintly striate. Pulvilli medium, adpressed. Parempodia flap-like, truncate. Trichobothria 10:7, trichomae weak. Bothria recessed.

Measurements: Total body 1. 3.80, 4.08. Head w. 0.98 n=1, 1. 0.84 n=1, vertex 0.55 n=1. Antenna segment 10.40, 0.42, II 1.48, 1.30. Rostrum total 1. 1.34, 1.36, IV 0.34, 0.34. Pronotum a.w. 0.90, 0.90, p.w. 1.28, 1.30, 1. 0.60, 0.58. Wing pad 0.76, 0.66. Metafemur 1. 1.42, 1.52, w. 0.40, 0.38. Metatibia 1. 1.92, 2.06. Metatarsus I 0.19, 0.20, II 0.56, 0.57. DAGO 0.09, 0.09. n=2 except where shown.

Material examined: 1 ex., Surrey, Bookham, 21.VI.55, EWG. 1 ex., Berks, Silwood Park, 27.VII.60, TRES.

Remarks: The host plant was not recorded for the above speci- mens, but it was very probably Galium palustre. The general colour of the nymphs was recorded by Southwood and Leston (1959). - 396 -

Polymerus unifasciatus (Fabricius) (S. Poeciloscytus) Figs. 19:170,171; 35:4; 61

Dark green. Meso- and meta-nota with internal margins and apices infuscate. Femora, tibiae with brown tinges. Tarsi, rostral apex infuscate. Claws black. Broad. Dorsal body hairs dense, black, stout, simple (max.l. 0.10). Ventral body hairs pale, slender, simple (max.l. 0.12). Tibiae with stout, striate, black spines. Cuticular sculpturing in form of spinules over entire body surface, except proximal dorsal half of femora. DAGO double, anterior bar sinuate, black patch anteriad and posteriad. Claws thick basally, striate. Pulvilli medium, rugulose. Parempodia large, flap-like, truncate, striate. Trichobothria 9:7, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.00-4.40 (4.14). Head w. 0.96- 1.04 (1.00), 1. 0.70-0.80 (0.77), vertex 0.50-0.54 (0.52). Antenna total 1. 3.27-3.45 (3.34) n=5, 10.36-0.40 (0.38)n=5, II 1.16-1.28 (1.22) n=5, III 0.78-0.90 (0.86) n=5, IV 0.87- 0.90 (0.88 n=5. Rostrum total 1. 1.30-1.46 (1.37), IV 0.34- 0.39 (0.36). Pronotum a.w. 0.84-1.00 (0.93), p.w. 1.20-1.38 (1.27), 1. 0.56-0.66 (0.61). Wing pad 0.60-0.70 (0.66). Metafemur 1. 1.44-1.63 (1.53), w. 0.40-0.46 (0.43). Meta- tibia 1. 1.88-2.14 (2.02). Metatarsus I 0.18-0.21 (0.19), II 0.50-0.56 (0.53). n=6 except where shown.

Material examined: 3 ex., Oxon, AstonRowant, 24.VII.65,GEW. 13 ex., Perths, Rannoch School, 24.VI.78, Galium saxatile, GMcG.

Remarks: Butler (1923) gave a good description of the nymphs of this species, although the•'clearly recognisable keel' mentioned by him is not evident. - 397 - Polymerus nigritus (Fallen) (S. Polymerus s.s.) Figs. 19:168,169; 35:6; 61

Brownish-black. Head, nota brownish-black, with well-marked pale ecdysial lines. Abdomen tinged red, terminal segment infuscate. Claws black. Broadly oval. Pronotum with posterior angles rounded. Dorsal body hairs stout, slender, curved, simple, black (max.l. 0.11). Ventral body hairs slender, simple, pale (max..1. 0.12). Femora with stout, black hairs. Tibiae with stout,striate spines. Cuticular sculpturing in form of dark spinules. DAGO slit-like, dark, anterior bar heavily sclerotised, straight and with black patch anteriad and posteriad. Claws striate, stout basally. Pulvilli medium, rugulose, adpressed. Parempodia flap-like, truncate. Tricho- bothria 10:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.04-4.72 (4.35). Head w. 0.90- 1.02 (0.97), 1. 0.76-0.92 (0.83), vertex 0.46-0.58 (0.54). Antenna total 1. 2.90-3.26 (3.12), I 0.38-0.42 (0.40), II 1.10 -1.18 (1.14), III 0.68-0.80 (0.75), IV 0.74-0.87 (0.82). Rostrum total 1. 1.26-1.56 (1.45), I 0.40-0.54 (0.45), II 0.36-0.39 (0.37), III 0.24-0.28 (0.26), IV 0.34-0.38 (0.36). Pronotum a.w. 0.78 -0.96 (0.88), p.w. 1.22-1.40 (1.33), 1. 0.56-0.64 (0.61). Wing pad 0.62-0.80 (0.69) n=9. Metafemur 1. 1.32-1.48 (1.42), w. 0.38-0.56 (0.43). Metatibia 1. 1.80-2.20 (197). Metatarsus 10.20-0.21 (0.20), II 0.50-0.56 (0.53). DAGO 0.10 (0.10) n=5. n=10 except where shown.

Material examined: 2 ex., Bucks, Slough, -.VI.53, GEW. 5 ex., Mddsx, Hounslow, -.VI.54, GEW. 11 ex., Surrey, Bookham, 4.VI.54, Galium aparine, TRES. 1 ex., Oxon, Hartswood, 9.VII.56, TRES.

Remarks: This species, described by Butler (1923), has also been taken on Galium verum, G. boreale and Stachys sylvatica. There is a strong resemblance to Orthocephalus nymphs. However, the stout, squat shape of rostral segments III and IV and the broad oval DAGO of the latter are sufficient to distinguish the two species. - 398 -

Charagochilus gyllenhali (Fallen) Figs. 19:172,173; 35:7

Green. Wing pads light brown with posterior margins strong- ly infuscate. Small, squat. Dorsal body hairs dark, simple (max.l . 0.09) . Ventral body hairs slender (max.l . 0.10) . Tibiae with pale,striate spines. DAGO double, anterior bar pale, slightly sinuate. Claws striate. Pulvilli small to medium. Parempodia large, flap-like. Trichobothria 9:7, trichomae compact. Bothria recessed.

Measurements: Total body 1. 3.00-3.48 (3.24). Head w. 0.80- 0.86 (0.83), 1. 0.60-0.70 (0.63), vertex 0.40-0.54 (0.46). Antenna total 1. 1.90-2.11 (2.00), I 0.28-0.34 (0.30), II 0.70-0.80 (0.73), III 0.38-0.46 (0.41), IV 0.50-0.58 (0.55). Rostrum total 1. 1.06-1.23 (1.14), I 0.30-0.35 (0.31), II 0.28-0.36 (0.32), III 0.19-0.20 (0.20), IV 0.28-0.30 (0.29). Pronotum a.w 0.77-0.82 (0.80), p.w. 1.08-1.18 (1.14), 1. 0.54-0.62 (0.58). Wing pad 0.45-0.58 (0.51). Metafemur 1. 0.94-1.01 (0.98 ), w. 0.30-0.34 (0.32) . Metatibia 1. 1.24- 1.36 (1.28). Metatarsusl 0.14-0.17 (0.15), II 0.37-0.40 (0.39). DAGO 0.01-0.02 (0.01) n=4. n=10 except where shown.

Material examined: 14 ex., Herts, Harpenden, 26.VII.53, Galium sp., TRES.

Remarks: Kullenberg (1944) gave a colour sketch of a Vth instar nymph. The ratio of metatibia length/head width for this species is 1.46-1.70 (1.55) n=10. - 399 -

Charagochilus weberi Wagner, 1953

Green. Wing pads light brown, becoming darker apically. Dorsal body hairs dark, simple (max.l. 0.11). Ventral body hairs pale, simple, slender.(max.l. 0.13). DAGO double, anterior bar slightly sclerotised, sinuate. Claws striate. Pulvilli small to medium. Parempodia large, flap-like. Trichobothria 8:7?5 trichomae present. Bothria recessed.

Measurements: Total body 1. 3.48, 3.60. Head w. 0.78, 0.80, 1. 0.60, 0.73, vertex 0.39,0.40. Antenna total 1. 2.58, 2.63, I 0.36,0.38, II 0.97,1.00, III 0.52, IV 0.70, 0.76. Rostrum total 1. 1.22,1.23, IV 0.32. Pronotum a.w. 0.68, 0.725 p.w. 1.09, 1.10, 1. 0.54;0.56. Wing pad 0.52, 0.56. Metafemur 1. 1.10,1.14, w. 0.27. Metatibia 1. 1.58,1.60. Metatarsus 10.16 n=1, II 0.42 n=1. n=2 except where shown.

Material examined: 2 ex., Hants, Pamber Forest, 25.1X.655 , GEW.

Remarks: The occurrence and biology of this species in Britain was described byWoodroffe (1966). The ratio of metatibial length/head width for this species is approximately 2. - 400 - Dichrooscytus rufipennis (Fallen) Fig. 19:174,175

Green. Eye facets dark red. Meso- and meta-nota with bright red markings, becoming apically black in late Vth instar. Tarsi, rostral apex infuscate. Abdominal segments with faint transverse banding. Rostrum extending beyond meta- thoracic coxae, joint between segments II, III very faint. Dorsal body hairs short, black, simple, straight or slightly curved (max.l. 0.08). Ventral body hairs pale, straight, slender, simple (max.I. 0.09). Tibiae with stout, black, striate spines, shorter than tibial diameter. DAGO not visible. Claws striate. Pulvilli very small . Parempodia flap-like. Trichobothria 8:6, trichomae absent. Bothria flush.

Measurements: Total body 1. 4.70-5.40 (4.92). Head w. 1.08 -1.14 (1.12), 1. 0.94-1.10 (0.98), vertex 0.51-0.56 (0.54). Antenna total 1. 3.28-3.78 (3.59) n=7, I 0.40-0.44 (0.42), II 1.42-1.48 (1.45), III 0.90-1.02 (0.96) n=7, IV 0.74-0.84 (0.79). Rostrum total 1. 2.20-2.48 (2.37) n=8, IV 0.65- 0.68 (0.66). Pronotum a.w. 0.93-1.06 (1.01) n=8, p.w. 1.20- 1.30 (1.25) n=8, 1. 0.57-0.65 (0.60). Wing pad 0.66-0.80 (0.75). Metafemur 1. 1.46-1.54 (1.51), w. 0.36-0.40 (0.39). Metatibia 1. 2.06-2.20 (2.12). Metatarsus 10.18-0.20 (0.19), II 0.58-0.62 (0.61). n=9 except where shown.

Material examined: 14ex., Berks, Silwood Park, 23.VI.77, Pinus sylvestris, GMcG. 10ex., Perths, Rannoch School, 24-25.VI.78, P. sylvestris,GMcG.

Remarks: The red colouration on the nymphal wing pads is complementary to the adult colouration, the entire hemi- lytra, except the membrane, being suffused with red. The metathoracic femoral trichs are weak and there are some setae about which the distinction between hair and trich is not clear. There is some variation in trichobothrial number in this species. Butler (1923) described the Vth instar nymph. The genus is unusual in apparently having no DAGO. - 401 - Dichrooscytus valesianus (Meyer-DUO

Claws, rostral apex, apical portion of tarsal segment II infuscate. Dorsal body hairs short, black, simple, slightly curved (max.l. 0.04). Ventral body hairs pale, long, simple. DAGO not visible. Claws striate. Pulvilli small. Parem- podia flap-like. Trichobothria 7:6, trichomae absent. Bothria flush.

Measurements: Total body 1. 3.24-3.36 (3.31) n=3. Head we 0.80-0.86 (0.84) n=4, 1. 0.64-0.70 (0.66) n=3, vertex 0.38, 0.40 n=2. Antenna total 1. 2.60, 2.74 n=2, I 0.32-0.34 (0.33) n=4, II 0.94-1.00 (0.96) n=4, III 0.72-0.80 (0.75) n=3, IV 0.60, 0.60 n=2. Rostrum total 1. 1.24-1.34 (1.28) n=4, IV 0.30-0.40 (0.38) n=4. Pronotum a.w. 0.73-0.76 (0.74) n=3, p.w. 0.92-0.99 (0.96) n=3, 1. 0.40-0.46 (0.44) n=3. Wing pad 0.47-0.59 (0.52) n=4. Metafemur 1. 1.04-1.12 (1.07) n=3, w. 0.30 (0.30) n=3. Metatibia 1. 1.46-1.54 (1.49) n=4. Metatarsus I 0.14 (0.14) n=4, II 0.46-0.48 (0.47) n=4.

Material examined: 2 ex., Oxon, Baldhill, 17.VI.60, GEW. 4 ex., Surrey, Riddlesdown, 12.VI.58, EWG.

Remarks: The colour of the nymphs is probably green with red tinges on the abdomen, and the host plant is Juniperis communis (Southwood & Leston, 1959). - 402 - Miris striatus (L.) Figs. 19:176,177; 36:2

Dark reddish-brown to black. Antennae brown, segment I light brown. Head, nota dark reddish-brown to black, with clearly marked ochreous ecdysial lines. Abdominal segments I, II with posterior margins yellow, other segments with posterior margins less strongly brown and with white flashes laterally. Femora, tibiae reddish-brown. Tarsi, claws black. Large, elongate. Antennae long. Lateral margins of wing pads sub- parallel. Abdomen sometimes swollen. Legs slender. Dorsal body hairs short, stout, dark, blunt (max.l. 0.08). Ventral body hairs long, simple. pale (max.l. 0.14). Tibiae with stout, short, black, striate spines. DAGO double, sclerotis- ed, anterior bar slightly sinuate. Claws stout, black, striate. Pulvilli medium to large, rugulose. Parempodia flap-like, strongly striate. Trichobothria 9:8, trichs short, trichomae absent. Bothria flush.

Measurements: Total body 1. 6.83-8.16 (7.59). Head wo 1.28- 1.36 (1.32), 1. 1.12-1.40 (1.32), vertex 0.40-0.60 (0.53). Antenna total 1. 6.91-7.08 (7.17), 10.92-1.04 (0.96), II 2.36-2.76 (2.48), III 1.76-1.96 (1.85), IV 1.80-1.96 (1.84). Rostrum total 1. 2.36-2.42 (2.44), I 0.56-0.72 (0.63), II 0.60-0.64 (0.61), III 0.48-0.62 (0.53), IV 0.64-0.72 (0.67). Pronotum a.w. 1.20-1.40 (1.32), p.w. 1.44-1.68 (1.59), 1. 1.04-1.16 (1.09). Wing pad 0.96-1.20 (1.13). Metafemur 1. 2.24-2.88 (2.69), w. 0.48-0.56 (0.52). Metatibia 1. 3.88- 4.40 (4.14). Metatarsus 10.32-0.36 (0.34), II 0.72-0.80 (0.77). DAGO 0.11 (0.11). n=8 in all cases.

Material examined: 2 ex., Lancs, Great Tower, 1.VI.54, Quercus, TRES. 2 ex., Lancs,.Eggerslack, 10.VI.54, Quercus, TRES. 2 ex., Bucks, Slough, 15.VI.54, Quercus, TRES. 3 ex., Berks, Silwood Park, 11.VI.57, Quercus, TRES. 2 ex., Oxon, Aston Rowant, 23.V.65, GEW. 1 ex., Dorset, Milton Abbas, 4.VI.78, Crataegus sp., GMcG.

Remarks: The size and colouration of this species, which was described by Butler (1923) and Kullenberg (1944), make it readily recognisable. Southwood and Leston (1959) gave - 403 - a list of prey items taken and the trees on which the species is commonly found. Waloff and Southwood (1960) included it in a key to the mirid nymphs occurring on broom. The trichobothria are hard to see, being difficult to distinguish from setae. It is not possible to say whether this genus is primitive in having very indistinct, short trichobothria with absent trichomae, or secondarily specialised for some reason. - 404 - Calocoris quadripunctatus (Villers) (S. ) Figs. 2c; 7b; 18a; 19:178,179; 36:4; 77

Yellow, reddish-brown and brown. Antennae brown with segment IV red. Head with 3 longitudinal wine red stripes from anterior margin to line with posterior margins of eyes. Eyes red . Pronotum pale yellow, brown patches symmetrical about body midline. Wing pads yellow to olive-green with brown mottling, patches and 3 longitudinal brown stripes. Abdomen yellow or yellowish-white with transverse red band- ing and mottling, occasionally dark reddish-brown. Femora, tarsi brown. Tibiae light brown with base and apex fuscous. Dorsal body hairs stout, black, blunt (max.l. 0.04), occasionally long, bifurcate on dorsal surface of terminal two abdominal segments (max.l. 0.05). Ventral body hairs pale, slender, simple (max.l. 0.11). Head with long hairs (max.l. 0.08). DAGO double, anterior bar slightly sinuate, sclerotised, with pigmented patchfosteriad. Claws thick basally, weakly striate. Pulvilli large, rugulose, detached apically. Parempodia flap-like, narrow, striate, truncate. Trichobothria 7:6, trichomae absent. Bothria slightly tuberculate.

Measurements: Total body 1. 5.24-6.41 (5.68). Head w. 1.02 -1.11 (1.07), 1. 0.90-1.00 (0.95) n=7, vertex 0.52-0.60 (0.57) n=8. Antenna total 1. 4.10-4.64 (4.48), I 0.74-0.86 (0.83), II 1.60-1.94 (1.81), III 1.10-1.30 (1.21), IV 0.60- 0.68 (0.64). Rostrum total 1. 2.00-2.26 (2.12) n=8, IV 0.57-0.62 (0.60) n=8. Pronotum a.w. 1.00-1.10 (1.06) n=8, p.w. 1.46-1.50 (1.49) n=7, 1. 0.70-0.80 (0.77) n=7. Wing pad 0.80-0.90 (0.85). Metafemur 1. 1.80-2.10 (2.02), w. 0.40-0.48 (0.44). Metatibia 1. 2.76-3.20 (3.04). Metatarsus 10.20-0.22 (0.21), II 0.49-0.58 (0.54) n=8. n=9 except where shown.

Material examined: 8ex., Westm?,Craggie Wood, 1.VI.54, Quercus sp., TRES. 2ex., Westm, Sizergh, 3.VI.54, Quercus, TRES. 1 ex., Surrey, Chobham Common, -.VI.58, Quercus,GEW. 2ex., Berks, Windsor, 21.V.61, Quercus sp., GEW. 1 ex., Dumfs, Bodesbeck, 1.VI.77, Quercus sp., GMcG. -405 -

Figure 77 - 406 -

Remarks: The distinctive patterning of this nymph makes it easily recognisable. The adults may be confused with those of Miris striatus, although the latter are much larger. Young nymphs have olive-green nota and legs and orangish-red abdomens. - 407 - Calocoris stysi Wagner, 1968 (S. Lophyromiris) sexguttatus var. insularis Reuter, 1896 nec insularis Horvath, 1879 Figs. 19:180,181; 36:5

Purplish-brown. Base of antennal segment I, segment IV entirely red, rest of antennae light brown. Head dark reddish- brown, shiny with pale red ecdysial lines. Pronotum shiny with calli large, dark brown, lateral anterior margins white. Wing pads shiny, brown with lateral margins and medial stripe white. Abdomen purplish-red or brown with posterior margins of segments ochreous, terminal two segments brown, shiny. Coxae, trochanters white. Femora, tibiae light brown. Tarsal segment I, base and apex of segment II and claw entirely infuscate. Head long, tapering behind eyes. Pronotum with anterior and posterior margins slightly indented, angles rounded. Meso- and meta-notum broader than pronotum. Abdomen occasionally swollen. Dorsal body hairs black, spinous, simple (max.l. 0.06). Ventral body hairs long, slender, simple (max.l. 0.12). Abdominal spiracular openings sclerot- ised, slightly expanded (diameter 0.03). DAGO double, anter- ior bar heavily sclerotised, sinuate, with small black patch anteriad and posteriad. Claws thick basally, lightly' striate. Pulvilli medium to large. Parempodia flap-like, striate. Trichobothria 7:5, trichomae weak. Bothria flush.

Measurements: Total body 1. 5.16-5.80 (5.53). Head w. 1.00- 1.08 (1.04) n=8, 1. 0.97-1.16 (1.05) n=8, vertex 0.39- 0.44 (0.41) n=8. Antenna total 1. 5.46-5.80 (5.64) n=7, I 0.76 -0.82 (0.80), II 1.76-1.94 (1.83), III 1.44-1.52 (1.49) n=7, IV 1.50-1.58 (1.52) n=7. Rostrum total 1. 2.18-2.32 (2.25) n=8, IV 0.62-0.66 (0.64). Pronotum a.w. 0.90-1.02 (0.95) n=8, p.w. 1.22-1.34 (1,25) n=8, 1. 0.64-0.76 (0.70) n=8. Wing pad 0.79-0.94 (0.88). Metafemur 1. 2.04-2.22 (2.15), w. 0.37-0.42 (0.39). Metatibia 1. 3.00-3.21 (3.14). Metatarsus I 0.20 (0.20), II 0.57-0.59 (0.58). DAGO 0.14 (0.14) n=5. n=9 except where shown.

Material examined:.12ex., Herts, Harpenden, 28.VI.55, Urtica dioica, TRES. 6ex., Som, Ubley, 4.VI.77, GWW. - 408 - Remarks: Butler (1923) described the nymphs and gave a list of host plants. However, Urtica dioica appears to be the commonest host. Southwood and Scudder (1956b)described the nymph and gave a drawing of a IVth instar nymph. The long antennae and the shiny, dark purplish-brown or red appearance make the nymphs of this species readily identi- fiable. - 409 - Calocoris alpestris (Meyer-DUO (S. Calocoris s.s.) Figs. 19:182,183; 36:6

Green with brown markings. Base of antennal segment I, apex of segments II, III and segment IV entirely brown. Head, pronotum largely green with faint brown markings on posterior margins. Wing pads green with brown markings on posterior margins, apices, anterio-lateral angles. Mesonotum with large pale green 'heart-shaped' patch. Femoral apices, basal and apical one-third of tibiae brown. tarsi, claws, rostral segment IV brown. Dorsal body hairs black, short, slightly curved, simple (max.l. 0.09). Ventral body hairs pale, long, simple (max.l. 0.11). Abdominal spiracular openings slightly expanded (diameter 0.028). DAGO double, anterior bar sclerotised, sinuate, with small dark patch anteriad and posteriad. Claws curved, lightly striate. Pulvilli medium to large. Parempodia flap-like, striate. Trichobothria 9:7 trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 6.08-6.66 (6.39). Head w. 1.12- 1.24 (1.19), 1. 1.00-1.06 (1.01), vertex 0.55-0.60 (0.58) n=3. Antenna total 1. 6.82, 6.92 n=2, I 0.88 -0.98 (0.92), II 2.16-2.40 (2.29), III 1.58-1.68(1.64)n=3, III 1.88,1.90 n=2. Rostrum total 1. 2.44-2.52 (2.48), IV 0.68-0.72 (0.69). Pronotum a.w. 1.00-1.14 (1.08), p.w. 1.52-1.68 (1.62), 1. 0.88-0.92 (0.89). Wing pad 1.12-1.22 (1.16). Metafemur 1. 2.56-2.96 (2.73), w. 0.44-0.50 (0.47) . Metatibia 1. 3.64- ' 3.80 (3.74). Metatarsus I 0.23-0.24 (0.24), II 0.76-0.80 (0.78). n=4 except where shown.

Material examined: 2 ex., Lancs, Merlewood, 21.V.54, Urtica dioica, TRES. 2 ex., Som, Norton St Philip, 29-300V.55 TRES.

Remarks: Southwood and Scudder (1956b)described and illus- trated the nymphs and included this species in a key to the immature stages of Heteroptera occurring on Urtica dioica. - 410 -

Calocoris norvegicus (Gmelin) (S. Calocoris s.s.) Figs. 19:184,185; 36:7

Green with occasional red tinges on abdomen and nota. Tarsi, claws, rostral tip infuscate. Dorsal body hairs fairly dense, black, slightly curved, striate, simple (max.1.0.11). Ventral body hairs paler, straight, simple (max.l. 0.11). Abdominal spiracularcpenings slightly expanded (diameter 0.02). DAGO double with anterior bar sinuate. Claws thick basally, lightly striate. Pulvilli medium. Parempodia flap-like, striate. Trichobothria 10:6, trichomae present. Bothria slightly recessed.

Measurements: Total body 1. 5.41 6.49 (5.79). Head w. 1.10- 1.22 (1.15), 1.0.90-1.06 (1.01), vertex 0.54-0.60 (0.59). Antenna total 1. 4.46-4.86 (4.73), I 0.64-0.70 (0.67), II 1.70-1.90 (1.83), III 1.10-1.20 (1.15), IV 1.02-1.12 (1.08). Rostrum total 1. 2.10-2.30 (2.21), I 0.58-0.66 (0.61), II 0.50-0.62 (0.57), III 0.38-0.44 (0.41), IV 0.60-0.64 (0.61). Pronotunr a.w. 0.98-1.16 (1.19), p.w. 1.40-1.59 (1.47), 1. 0.74-0.92 (0.80) . Wing pad 0.68-0.94 (0.84) . Netafemur 1. 2.00-2.30 (2.12), w.0.46 -0.56 (0.52). Metatibia 1. 2.84- 3.12 (2.99). Metatarsus I 0.19-0.26 (0.23), 1I0.54-0.74 (0.69). DAGO 0.105 (0.105) n=4. n=10 except where shown.

Material examined: 2 ex., Oxon, Hartswood, 24.VI.50,TRES. 14 ex., Herts, Great Field, 2.VII.53, TRES.

Remarks: Butler (1923) described the nymphs and recorded a list of host plants. Southwood and Scudder (1956b)illustr- ated the Vth instar nymph and included this species in a key to the Heteroptera associated with Urtica dioica on which it is commonly found. - 411 -

Calocoris roseomaculatus (DeGeer) (S. calocoris s.s.) Figs. 19:186,187; 36:8

Reddish-brown. Head, nota with thin ochreous ecdysial lines. Abdomen faintly transversely banded, segments I, II ochreous. Tarsi, claws black. Oval, slightly shiny. Pro- notum with posterior half slightly flared. Dorsal body hairs black, short, slightly curved, simple (max.l. 0.05), occasionally long, black,stout, bifurcate or serrate (max. 1. 0.08). Ventral body hairs long, pale, simple (max.l. 0.09). Femora, tibiae with stout, black, striate spines. DAGO double, margins fuscous with anterior bar sclerotised, sinuate. Claws striate. Pulvilli medium. Parempodia flap- like. Trichobothria 10:6, trichomae present. Bothria slightly recessed.

Measurements: Total body 1. 5.83-6.33 (6.05). Head w. 1.20- 1.26 (1.22), 1. 1.10-1.18 (1.14), vertex 0.48-0.56 (0.54). Antenna total 1. 5.77-6.20 (6.06), I 0.70-0.76 (0.74), II 2.09-2.30 (2.19), III 1.58-1.66 (1.62), IV 1.40-1.52 (1.48). Rostrum total 1. 2.56-2.84 (2.74), IV 0.72-0.79 (0.76). Pronotum a.w. 1.14-1.20 (1.18), p.w. 1.52-1.68 (1.58), 1. 0.87-1.04 (0.94). Wing pad 0.70-1.00 (0.91). Metafemur 1. 2.09-2.29 (2.20), w. 0.50-0.58 (0.55). Metatibia 1. 3.12- 3.36 (3.28). Metatarsus I 0.30-0.34 (0.32), II 0.65-0.70 (0.68). DAGO 0.15 (0.15). n=5 in all cases.

Material examined: 1 ex., Bucks, Slough, -.VI.53, GEW. 2 ex., Surrey, Farthing Downs Coulsdon, 3.VII.54, TRES. 3 ex., Suffolk, East Bergholt, 11.VII.54, TRES. 1 ex., Surrey, Witley Common, -.VI.58, GEW. 1 ex., Inverness, Aviemore, 26.VI.67, GEW.

Remarks: The host plants mentioned for this species are Sanguisorba minor on chalk slopes and Lotus corniculatus, Trifolium sp. and Ononis sp. in other areas (Southwood & Leston, 1959). - 412 -

Adelphocoris lineolatus (Goeze) Figs. 7c; 19:188,189; 37:1; 61; P1 16

Dark green. Antennae brownish-black. Head, nota tinged brown. Femora, tibiae heavily spotted with small brown spots. Tarsi brownish-black. Claws black. Dorsal body hairs stout, black, striate, bifurcate or blunt arising from brown or black spots (max.l. 0.146) or small, black, striate, simple or bifurcate not arising from brown or black spots (max.l. 0.087). Ventral body hairs pale, simple, not arising from pigmented spots (max.l. 0.17). Femora, tibiae with black, stout spines arising from pigmented spots. Abdominal spiracular opening slightly expanded (diameter 0.026). DAGO single, constricted medially, anterior bar slightly curved, with pigmented area anteriad and posteriad. Claws striate. Pulvilli medium, adpressed. Parempodia flap-like, striate. Tr5.chobothria 9 or 10: 7, trichomae present, often associat- ed with small pigmented spot. Bothria flush.

Measurements: Total body 1. 5.66-6.33 (6.10). Head w. 1.12- 1.20 (1.15), 1. 1.00-1.18 (1.06), vertex 0.50-0.60 (0.57). Antenna total 1. 5.29-5.92 (5.67), I 0.74-0.88 (0.80), II 1.96-2.18 (2.01), III 1.56-1.80 (1.67), IV 1.10-1.24 (1.19). Rostrum total 1. 2.42-2.70 (2.54), I 0.58-0.76 (0.70), II 0.60-0.70 (0.65), III 0.40-0.50 (0.46), IV 0.68-0.78 (0.72). Pronotum a.w. 1.00-1.10 (1.07), p.w. 1.32-1.54 (1.42), 1. 0.66-0.86 (0.76). Wing pad 0.78-1.18 (0.94) n=9. Metafemur 1. 2.30-2.72 (2.52), w. 0.46-0.57 (0.53). Metatibia 1. 3.24- 3.88 (3.52). Metatarsus I 0.20-0.26 (0.24), II 0.65-0.80 (0.72). n=10 except where shown.

Material examined: 11 ex., Beds, Dunstable Downs, 5.VIII.54, Ononis sp., TRES. 1 ex., Hants, Pamber Forest, 6.VIII.59, GEW. 2 ex., Surrey, Witley Common, 3.VIII.65, GEW.

Remarks: Southwood and Leston (1959) gave a list of host plants and Butler (1923) and Puchkov and Puchkova (1956) described the nymphs. - 413 -

Adelphocoris seticornis (Fabricius) Fig. 37:2

Material examined: 2 IVth ex., Devon, Braunton, 1.VII.59, GEW. 1 IVth ex., Isle of Wight, Totland Bay, 2.VII.60, GEW.

Remarks: The head and nota are slightly shiny. The spotting on the nymphs of this species is not nearly as marked as in the preceding species. Butler (1923) and Southwood and Leston (1959) gave lists of possible host plants which included Lathyrus pratensis, Lotus uliginosus and cracca. Butler (1923) gave the following description: ' Head black; pronotum brown with central furrow; abdomen purplish brown with a basal yellow band; wing pads almost black, with a yellowish lateral streak at the shoulder; femora reddish yellow with darker spots, tibiae testaceous with black apex and black spinous hairs; tarsi black wish basal half of second joint yellow; antennae with basal joint yellowish, second reddish brown, darker at apex, third and fourth black, the former with base and the latter with the extreme apex reddish yellow.' Puchkov and Puchkova (1956) described the nymphs. - 414 - Adelphocoris ticinensis (Meyer-DUr) Fig. 37:3

Purplish-brown and green. Antennal segment I red, segment II and basal half of segment III reddish-green, apical half of segment III and all segment IV dark purplish-red. Head purplish-brown with broad pale markings between eyes, extending longitudinally, joining at posterior margin of head. Nota largely green. Abdomen purplish-brown (early instars), becoming green tinged with purple. Femora mottled purplish-brown, tibiae less mottled, light brown. Apical half of tarsal segment II and claws entirely dark fuscous to black. Dorsal body hairs black, spinous, not arising from pigmented spots (max.l. 0.08), and on abdomen, head and anterior pronotal angles mostly simple or blunt (max.l. 0.13) . Ventral body hairs pale, slender, simple (max.l. 0.10). Antennal segment I with 4 black spines. Femora,•tibiae with stout, black spines. DAGO single, darkly pigmented with anterior bar slightly curved. Claws striate. Pulvilli medium, adpressed. Parempodia flap-like, striate. Tricho- bothria 8:7, trichs fairly long, trichomae strong. Bothria recessed.

Measurements: Total body 1. 4.00-5.56 (4.70) n=5. Head w. 1.00-1.04 (1.02) n=5, 1. 0.82, 0.84 n=2, vertex 0.47-0.52 (0.50) n=5. Antenna total 1. 4.70-5.38 (5.00) n=4, I 0.56- 0.74 (0.64) n=5, II 1.60-1.88 (1.73) n=5, III 1.50-1.65 (1.5$) n=5, IV 1.04-1.12 (1.10) n=4. Rostrum total 1. 3.20, 3.20 n=2, IV 0.60, 0.60 n=2. Pronotum a.w. 1.00, 1.06 n=2, p.w. 1.20, 1.30 n=2, 1. 0.70, 0.74 n=2. Wing pad 0.54 (0, 0.92 (d). Metafemur 1. 2.08, 2.09 n=2, w. 0.50, 0.54 n=2. Meta- tibia 1. 2.64-2.96 (2.73) n=5. Metatarsus 10.28, 0.29 n=2, II 0.60, 0.68 n=2. DAGO 0.11,.0.11 n=2.

Material examined: 1 ex., Hants, Fleet Pond, 2.VIII.58, GEW. 2 ex., Surrey, Virginia Water, -.VII.59, GEW. 2 ex., Berks, Silwood Park, 27.VII.60, TRES.

Remarks: Southwood and Leston (1959) gave a list of host plants including Lotus uliginosus and Lathyrus palustris. - 415 - Megacoelum infusum (Herrich-Schhffer) Figs. 7d; 19:190,191; 37:5; 78; Pls 8; 9

Brown and red with distinctive red markings. Antennal segment I pale brown, II pale brown becoming tinged with red apically, III light brown basally, red apically and IV bright red. Head light brown with white ecdysial lines bordered with red, and with red lines round eyes extending to antennal sockets and posteriad to anterior margin of pronotum. Pronotum brown with central white line bordered with red, running from anterior to posterior margins, pleura occasionally with longitudinal red markings. Meso- and meta-thoracic nota brown with central white line bordered with red continuing from pronotum. Wing pad apices brownish-black. Abdomen light brown or reddish-brown with lateral rows of faint brown spots, segment II with anterior half red. Femora reddish- brown, faintly spotted and marked. Tibiae light brown to brown with small black marks basally and with faint longi- tudinal red stripes. Tarsi with apical half infuscate. Dorsal body hairs dark, coronate or blunt, parallel-sided (max.l. 0.18). Ventral body hairs pale, thin,simple (max.l. 0.17). Tibial hairs black, spinous. Metathoracic tibial hairs up to twice tibial diameter. Femoral cuticular sculp- turing as irregular transverse rows of pimples. DAGO single, constricted medially, anterior bar slightly curved. Claws striate. Pulvilli medium -large,rugulose, detached apically. Parempodia large, flap-like, striate. Trichobothria 8:6, trichomae absent. Bothria tuberculate.

Measurements: Total body 1. 4.74-5.24 (4.91). Head w. 0.90- 1.04 (1.00), 1. 0.80-0.90 (0.86), vertex 0.44-0.49 (0.47). Antenna total 1.5.45-6.08 (5.85), I 0.85-0.96 (0.90), II 1.92-2.22 (2.08), III 1.60-1.80 (1.72), IV 1.10-1.21 (1.13) Rostrum total 1. 2.16-2.29 (2.24), IV 0.64-0.68 (0.66). Pronotum a.w. 0.86-1.00 (0.91), p.w. 1.18-1.34 (1.26), 1. 0.65-0.70 (0.68). Wing pad 0.70-0.84 (0.79). Metafemur 1. 2.12-2.32 (2.24), w. 0.36-0.40 (0.38). Metatibia 1. 3.16- 3.32 (3.20). Metatarsus I 0.21-0.22 (0.21), II 0.58-0.62 (0.60). n=7 in all cases.

Material examined: 4 ex., Ashvale, 3.VIII.53, Pinus sp.,GEW. - 416 - Figure 78

Megacoelum infusum- (Vth instar nymph) - 417 - 4 ex., Mddsx, Hounslow, 8.VIII.53, Quercus sp., GEW. 1 ex., Surrey, Oxshott, 28.VI.57, Pinus sp., DL. 1 ex., Hants, Fleet Pond, -.VIII.59, Pinus sp. GEW. 1 ex., Beds, Am pthill, 6.VIII060, Quercus sp., DL. 1 ex., Beds, Studham, 12.VIII.60, Quercus sp., DL. 5ex., London, Putney Heath, 30.VII.76, Quercus sp., GMcG. 4 ex., London, Putney Heath, 27.VII.77, Quercus sp., GMcG.

Remarks: On the continent Megacoelum infusum occurs on oak and the very closely related M. beckeri on pine, both these animals being predacious on small insects (Wagner & Weber, 1964). The long metatibial hairs used to distinguish the two species do not seem to separate the British Megacoelum adult specimens examined. Dissection of d' adults from pine and oak in Britain labelled as beckeri and infusum respectively revealed that the genitalia were of one con- figuration, namely the of M. infusum. It would appear that in the absence of competition from M. beckeri on pine M. infusum has expanded to fill the niche in Britain. The nymphs collected from pine and oak appeared to be identical. Butler (1923) gave nymphal descriptions for both species, but it is possible that they refer to M. infusum.

In early instars the femora are red and the tibiae are strong- ly longitudinally marked with thin red stripes. In addition there are lateral red lines on the pleura extending to the coxae. The antennae of this active nymph are very long and the nymph has difficulty in drawing the last segment through the prothoracic tibial combs. - 418 -

Stenotus binotatus (Fabricius) Figs. 19:192,193; 37:6

Olive green. Head, nota, legs dark olive green. Tarsi, claws, apices of wing pads and apical half of rostral segment IV infuscate. Antennae long. Rostrum reaching just past metathoracic coxae. Dorsal body hairs stout, black, faintly striate, simple (max.l. 0.08). Ventral body hairs long, slender, pale, simple (max.1. 0.10) . Meso- and meta- thoracic femora with two strong spines apically. DAGO single slit-like, anterior bar pale, slightly curved. Claws striate. Pulvilli small-medium. Parempodia flap-like, striate. Tricho- bothria 9:6, trichomae very weak. Bothria flush/tuberculate.

Measurements: Total body 1. 4.33-5.49 (5.36). Head w. 0.94- 1.04 (0.99), 1. 0.82-0.96 (0.86), vertex 0.43-0.54 (0.50). Antenna total 1. 3.68-4.14 (3.90), 10.50-0.58 (0.54), II 1.30-1.50 (1.41), III 1.02-1.18 (1.09), IV 0.78-0.92 (0.86). Rostrum total 1. 1.92-2.24 (2.13), IV 0.50-0.58 (0.55) . Pronotum a.w. 0.80-0.96 (0.90), p.w. 1.12-1.36 (1.23), 1. 0.56-0.68 (0.63) . Wing pad 0.62-0.88 (0.76) . Metafemur 1. 1.44-1.64 (1.58), w. 0.38-0.44 (0.41). Metatibia 1. 2.08- 2.43 (2.28). Metatarsus 10.26-0.30 (0.28), II 0.40-0.52 (0.48). n=10 in all cases.

Material examined: 14 ex., Herts, Harpenden, 8.VII.53,TRES.

Remarks: Butler (1923) gave a brief description of the nymphs of this species which appear to suffer heavy para- sitism by braconids (Southwood, unpublished). They can be found on the flower heads Dactylis glomerata in early instars and Alopecurus pratensis and Phletmi pratense in the last two nymphal instars and the adult stage (Southwood & Leston, 1959). - 419 - Miridius cuadrivirgatus (Costa) Fig. 19:194,195

Green with one median and two lateral longitudinal white stripes running the entire length of body. Pronotum with lateral and posterior margins largely green. Abdomen with two extra longitudinal white stripes laterally. Femora, tibiae green with metathoracic femora mottled brown. Apical half of tarsal segment II and claws entirely infuscate. Elongate. Head elongate. Antennae long. Rostrum long, reaching well past metathoracic coxae. Dorsal body hairs black, slender, simple (max.l. 0.33). Ventral body hairs pale, slender, simple. Antennae with dense pubescence of long, slender hairs. DAGO double, anterior bar sinuate, weakly sclerotised. Claws stout basally, weakly striate. Pulvilli medium, detached apically. Parempodia flap-like, striate. Trichobothria 10:7, trichomae absent. Bothria flush.

Measurements: Total body 1. 5.40-5.99 (5.71) n=5. Head w. 1.06 n=1, 1. 1.00, 1.10 n=2, vertex 0.55 n=1. Antenna total 1. 5.72-6.28 (5.98) n=5, I 1.04-1.12 (1.08) n=5, II 1.92-2.08 (1.99) n=5, III 1.52-1.72 (1.62) n=5, IV 1.16- 1.36 (1.29) n=5. Rostrum total 1. 3.32, 3.44 n=2, IV 0.72, 0.73 n=2. Pronotum a.w. 1.00, 1.08 n=2, p.w. 1.28, 1.40 n=2, 1. 0.66, 0.77 n=2. Wing pad 0.96, 1.00 n=2. Meta- femur 1. 2.64, 2.80 n=2, w. 0.58, 0.60 n=2. Metatibia 1. 3.52-3.88 (3.71) n=5. Metatarsus I 0.26, 0.28 n=2, II 0.63, 0.70 n=2. DAGO 0.08 n=2.

Material examined: 3 ex., Devon, Totnes, 9.VII.16, EAB. 2 ex., Devon, Wembury, 4.VII.60,DL.

Remarks: Butler (1923) described the distinctive nymphs of this species, which are associated with Hordeum murinum. stripes The green longitudinal/formed by the white longitudinal stripes are of the same width as the white stripes. The appearance of these nymphs is very reminiscent of those of the Stenodemini. However, they possess a high metafemoral trichobothrial number typical of the Mirini and flush bothria. - 420 -

Phytocoris tiliae (Fabricius) (S. Phytocoris s.s.) Figs. 19:196,197; 42e; P1 14

Pale greenish-brown. Antennal segment I reddish-brown, II and III with two broad brown bands, IV evenly pale brown. Head largely pale with reddish-brown mottling, lateral mar- gins dark brown to black posterior to eyes and around antennal sockets. Pronotal lateral margins dark brown to black, posterior half with brown mottling. Wing pads pale, mottled with light brown, apices black. Abdomen largely mottled with reddish-brown and white, with some reddish- brown spots laterally. Pro- and meso-thoracic femora pale mottled brown in broad bands, metathoracic femor with two or three broad,transverse,mottled brown bands. Tibiae with 3 broad,transverse,brown bands. Tarsi and claws light brown. Antennae long, body and legs elongate. Dorsal body hairs pale or black, very long, slender, simple. Antennal segment I with many long, simple hairs, held at right angles to segment. DAGO single, small, constricted medially, anterior bar slightly curved, with pigmented patch anteriad and posteriad. Claws striate. Pulvilli medium. Parempodia flap- like, striate. Trichobothria 8:7, trichomae very weak. Bothria slightly recessed.

Measurements: Total body 1. 4.08-4.50 (4.28) n=9. Head w. 0.80-0.88 (0.85) n=10, 1. 0.64-0.76 (0.70) n=10, vertex 0.38-0.44 (0.42) n=10. Antenna total 1. 4.56-5.02 (4.88) n=9, I 0.90-1.01 (0.96) n=10, II 1.52-1.70 (1.64) n=10, III 1.24-1.42 (1.33) n=9, IV 0.88-0.98 (0.94) n=9. Rostrum total 1. 2.00-2.10 (2.04) n=6, IV 0.52-0.56 (0.55) n=9. Pronotum a.w. 0.69-0.80 (0.75) n=8, p.w. 0.86-1.04 (0.98) n=8, 1. 0.49-0.54 (0.52) n=9. Wing pad 0.54-0.76 (0.65) n=10. Metafemur 1. 2.00-2.28 (2.20)'n=7, w. 0.41-0.45 (0.43) n=7. Metatibia 1.3.00-3.40 (3.27) n=7. Metatarsus I 0.17-0.20 (0.19) n=7, II 0.46-0.54 (0.50) n=7. DAGO 0.058 n=5.

Material examined: 10 ex., London, Hampstead Heath, 2.VZI.76, Betula sp., GMcG. 2 ex., Surrey, Richmond Park, 1O.VIII.76, Quercus sp., GMcG. 18 ex., London, Putney Heath, 27.VII.77, Quercus sp., GMcG. - 421 - Remarks: The form and colouration of the nymphs of this distinctive genus make them easily recognisable, but not so easily distinguishable from each other. P. tiliae is distinguished by the brown-black markings nn the lateral pronotal margins. The nymphs are very active and some spec- ies possess some slight saltatorial ability.

Because of a lack of accurately determined material, a key to this genus is not given, but will be the subject of a further study. - 422 - Capsus ater (L.) Figs. 2f; 3a; 7e,f; 18c; 19:198,199; 30:1

Dull parplish-red. Antennal segment I, II reddish-brown, III pale basally, light reddish-brown apically, IV reddish- brown. Head, nota dull brownish-red with white ecdysial lines. Abdomen becoming red with white transverse markings on posterior segmental margins, faint white spots all over. Femora with two faint mottled, broad, transverse bands apically. Tibiae with distal apices infuscate, tarsi,claws infuscate. Squat. Antennal segments Is II slightly incrassate, IV slender. Dorsal body hairs straight, simple or flabellate, ribbed, often serrate apically (max.l. 0.06). Femora, tibiae, antennae without flabellate hairs. DAGO double with anterior bar sinuate, sclerotised. Claws thick basally, faintly striate. Pulvilli small-medium. Parempodia flap-like, striate. Trichobothria 8:7, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 4.60-5.10 (4.93). Head w. 1.28-1.32 (1.30), 1. 0.90-1.00 (0.94), vertex 0.66-0.70 (0.68). Antenna total 1. 3.30-3.40 (3.35) n=3, I 0.50-0.56 (0.53), II 1.06-1.19 (1.13), III 0.60-0.70 (0.66) n=4, IV 0.92-0.99 (0.95) n=3. Rostrum total 1. 1.90-2.00 (1.96), IV 0.46-0.50 (0.47). Pronotum a.w. 1.18-1.28 (1.24), p.w. 1.54-1.60 (1.57), 1. 0.70-0.76 (0.72). Wing pad 0.59-0.68 (0.65). Metafemur 1. 1.36-1.58 (1.48), w. 0.42-0.47 (0.45). Metatibia 1. 1.90-2.09 (2.03). Metatarsus I 0.22-0.23 (0.22), II 0.50-0.56 (0.52). DAGO 0.10. n=5 in all cases.

Material examined: 4 ex., Herts., Royston, 7.VI.12, EAB. 1 ex., Bucks., Slough, -.VI. 53, GEW. 1 ex., Herts, Harpenden, 28.VI.55, TRES. 2•ex., Hants, Keyhaven, 5.VI.59, GEW. 6 ex., Perths, Kinloch Rannoch, 26.VI.78, GMcG. 2 ex., Dumfs, Bodesbeck Farm, 10.VII.78, MMM.

Remarks: The squat shape and flabellate hairs make the nymphs of this species readily identifiable. Southwood and Leston (1959) gave various grasses as the host plants, e.g. Lolium perenne and Agropyron repens, and made reference to the habit of the nymphs which are seldom taken by sweeping. - 423 -

The nymphs and adults are best taken by examining the bases of grasses in damp meadows. It is likely that the nymphs feed on the stolons and low parts of the grasses and that the flabellate hairs may be associated with the occupation of this niche. Various species of myrmecophilous root aphid possess spatulate and flabellate hairs on their dorsal surfaces exactly similar to those of C. ater. It has been suggested that these hairs may be associated with the retention of moisture or the protection of the animal from abrasion (Paul, 1977 unpublished). The similarities of the hairs and habits suggest that they share a causal link. The flabellate hairs appear to be longer and occur in great- er numbers on the dorsal surface of the head and pronotum, but it is by no means certain thatihey occur in sufficient numbers to afford any substantial protection from abrasion.

The nymphs captured in Dumfrieshire possessed a clear medial, whitish-ochreous band on each tibia and were not spotted on the abdomen. Examination of the adult d genitalia from the Dumfrieshire and Perthshire populations showed some morphological variation, but this has not been quantified. - 424 -

Pantilius tunicatus (Fabricius) Figs. 19:200,201; 38:2

Green with abdominal black spots. Antennae reddish-brown, segment II dark reddish-brown apically, III with apical half dark reddish-brown, IV entirely red. Head reddish-brown with short longitudinal black stripe between eyes and sparse small, black spots. Nota reddish-brown with sparse, small, black spots, thin black lines on lateral margin and thin, white ecdysial lines. Posterior margins of abdominal segments with small black spots arranged symmetrically about body mid- line. Apical half of tarsal segment II black. Rostrum reach- ing metathoracic coxae. Dorsal body hairs short, black, simple (max.l. 0.06). DAGO double, darkly pigmented, anterior bar sinuate, with pigmented patch anteriad and posteriad. Claws squat, striate. Pulvilli lobe-like. Parempodia flap- like, striate. Trichobothria 7:5, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1. 5.74, 6.08. Head w. 1.20, 1.24, 1. 0.90, 1.06, vertex 0.56, 0.63. Antenna total 1. 4.54, 5.28, I 0.90, 1.00, II 2.08, 2.44, III 0.94, 1.04, IV 0.62, 0.70. Rostrum total 1. 1.70, 1.80, IV 0.44, 0.48. Pronotum a.w. 1.10, 1.14, p.w. 1.68, 1.70, 1. 0.72;0.80. Wing pad 0.98, 1.14. Metafemur 1. 1.55, 1.66, w. 0.40, 0.41. Meta- tibia 1. 2.24, 2.56. Metatarsus I 0.22, 0.23, II 0.48, 0.51:. n=2 in all cases.

Material examined: 1 ex., (no data), Corylus avellana, TRES 1 ex., Surrey, Ockham, 15.IX.59, Betula sp., DL.

Remarks: Butler (1923) gave a description of the nymphs. - 425 - Acetroyis gimmerthali (Flor) Figs. 19:206,207; 38:7; 61

Green with longitudinal white stripe from posterior margin of head to apex of abdomen. Antennal segment IV with red tinge. Head with medial and two lateral,broad, brown stripes. Eyes whitish-ochreous. Pronotum with two longitudinal, brown stripes on either side of medial white stripe and two lateral longitudinal brown bands, lateral margins whitish-ochreous. Wing pads brown with lateral margins whitish-ochreous. Claws,apex of tarsal segment II black. Elongate, fusiform. Rostral tip reaching just past meso- thoracic coxae. Posterior margins of pronotum indented medi- ally. Dorsal body hairs thick, black, spinous, occurring mainly on the brown pigmented areas. DAGO narrow, double, anterior bar weak, curved. Claws smooth. Pulvilli very small, adpressed. Parempodia flap-like, lightly striate. Trichobothria 5:4, trichomae absent or very weak. Bothria flush.

Measurements: Total body 1. 5.00-5.83 (5.37). Head w. 0.80-0.94 (0.89), 1. 0.64-0.82 (0.76), vertex 0.52-0.66 (0.62). Antenna total 1. 3.22-3.50 (3.37), 10.58-0.68 (0.63), II 1.36-1.62 (1.50), III 0.70-0.78 (0.73), IV 0.50-0.52 (0.50). Rostrum total 1. 1.80-2.30 (2.09), IV 0.44-0.54 (0.50). Pronotum a.w. 0.80-1.00 (0.95), p.w. 1.20-1.42 (1.35), 1. 0.50-0.64 (0.61). Wing pad 0.74-0.82 (0.78). Metafemur 1. 1.12-1.36 (1.26), w. 0.30-0.36 (0.33). Meta- tibia 1. 1.62-1.88 (1.75). Metatarsus I 0.24-0.28 (0.27), II 0.42-0.46 (0.45). DAGO 0.092-0.115 (0.099). n = 10 in all cases.

Material examined: 4 ex., Berks, Silwood Park, 20.VIII.50, TRES. 9 ex., Surrey, Bookham, 4.VII.54, TRES. 4 ex.,Surrey, Witley Common, 14.VI.61, GEW. 23 ex., Berks, Silwood Park, 3.VII. 77, GMcG.

Remarks: Butler (1923) described the nymphs which occur on Arrhenatherum elatius and other grasses on commons, marshes and waste ground (Southwood & Leston, 1959). This species has the lowest trichobothrial number in the Stenodemini. - 426 - Stenodema calcaratum (Fallen) (S. Brachystira) Figs. 8b,c; 19:208,209; 38:8; 61

Brownish-green. Antennae light brown with reddish tinge. Head with lateral reddish-brown bands and medial white stripe continuing down nota. Abdomen green with pale yellowish- white long central stripe and margins with two longitudinal dull red stripes laterally near the connexivum. Femora with brown spots ventrally. Legs light brown. Claws, rostral tip and apex of tarsal segment II brownish-black. Elongate, fusi- form. Metafemora with one large spine distally on posterior margin. DAGO double, anterior bar sinuate. Claws slender, striate. Pulvilli small, detached apically. Parempodia flap-like. Trichobothria 7:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.66-6.41 (5.59). Head w.0.82- 0.96 (0.89), 1. 0.86-0.96 (0.90), vertex 0.42-0.58 (0.51). Antenna total 1. 4.04-4.70 (4.46), I 0.76-0.92 (0.84), II 1.62-1.96 (1.82), III 1.04-1.14 (1.06), IV 0.68-0.80 (0.74). Rostrum total 1. 1.92-2.16 (2.04), IV 0.46-0.48 (0.49). Pronotum a.w. 0.86-0.96 (0.91), p.w. 1.04-1.36 (1.15), 1. 0.63-0.76 (0.70). Wing pad 1.00-1.24 (1.10). Metafemur 1. 1.64-1.88 (1.69), w. 0.30-0.36 (0.33) . Metatibia 1. 1.78- 2.02 (1.99). Metatarsus I 0.30-0.38 (0.35), II 0.46-0.52 (0.50). DAGO 0.09-0.12 (0.10). n=9 in all cases.

Material examined: 8 ex., Herts, Harpenden, 24.VII.53, (bred), TRES. 2 ex., Herts., Harpenden, 30.VII.53, TRES. 3 ex., Devon, Braunton, -.IX.66, Phragmites sp., GEW. 1 ex., Berks, Silwood Park, 16.VII.76, GMcG. 2 ex., Dumfs, Bodesbeck Farm, 2.IX.76, Avena sativa, GMcG.

Remarks: The adults of this species have two metafemoral spines, the Vth instar nymphs one spine and the IVth instar nymphs a rudimentary spine. They have no aulpturing are slightly infuscate apically and appear to bear a primitive trichobothrium. Butler (1923), Ellis (1939) and Buczek (1959) gave details of biology and morphology. Southwood and Leston (1959) listed host plants including Agrostis tenuis and Alo9ecurus pratensis. - 427 -

Stenodema trispinosum Reuter (S. Brachystira) Figs. 19:210,211; 39:1; 61

Brown with reddish tinges. Head, nota with pale cream, medial ecdysial lines, becoming broad on abdomen, and bordered by two broad longitudinal light reddish-brown stripes. Nota and abdomen with red tinges, lateral margins pale. Abdomen with lateral rows of fuscous spots on connexivum. Tarsi, claws fuscous. Elongate, fusiform. Metathoracic femora with one large spine and one smaller spine distally on posterior margin. Dorsal body hairs short, black, spinous. DAGO double, narrow, anterior bar sinuate. Claws black, stout, striate. Pulvilli small. Parempodia flap-like. Trichobothria 6:5, trichomae present. Bothria recessed.

Measurements: Total body 1. 5.33-6.41 (5.72). Head w. 0.90- 0.96 (0.93), 1. 0.88-0.92 (0.91), vertex 0.50-0.54 (0.52) . Antenna total 1. 3.88-4.46 (4.23) n=9, I 0.74-0.80 (0.78), II 1.66-1.94 (1.76), III 0.86-1.06 (0.98), IV 0.64-0.78 (0.71)n=9. Rostrum total 1. 1.64-1.90 (1.80), IV 0.40-0.48 (0.45). Pronotum a.w. 0.94-1.04 (0.98), p.w. 1.22-1.36 (1.29), 1. 0.62-0.76 (0.71). Wing pad 1.02-1.14 (1.09). Metafemur 1. 1.60-1.80 (1.73), w. 0.38-0.42 (0.40). Metatibia 1. 1.92-2.10 (1.98). Metatarsus 10.32-0.36 (0.34), II 0.50- 0.54 (0.52). DAGO 0.10-0.12 (0.11). n=10 except where shown.

Material examined: 11 ex., Essex, Flatford, -.VIII, 49, Phragmites sp., TRES. 3 ex., Kent, Reculver, 15.IX.60, Phragmites sp., GEW.

Remarks: The adults of this species have three metafemoral spines, the Vth instar nymphs two spines,(Ellis, 1939). The larger nymphal spine is not as long or curved as that of S. calcaratum. - 428 -

Stenodema holsatum (Fabricius) (S. Stenodema s . s . ) Figs. 19:212,213; 39:2; 61

Dark green with yellow tinges. Antennal segment IV with reddish tinge. Head, nota with two lateral broad brown stripes and thin medial white stripe extending to wing pad apices. Nota, abdomen with lateral margins pale ochreous. Tarsi, claws, apex of rostrum black. Elongate, fusiform. Rostrum reaching beyond metathoracic coxae. Dorsal body hairs short, black, spinous. DAGO double, narrow, anterior bar sinuate, weakly sclerotised. Claws slender, striate. Pulvilli small. Parempodia flap-like, striate. Trichobothria 7:7, trichomae present. Bothria recessed.

Measurements: Total body 1. 6.23-6.66 (6.41). Head w. 0.96-1.06 (1.01), 1. 0.82-0.94 (0.89), vertex 0.50-0.60 (0.56). Antenna total 1. 4.40-5.26 (4.94), 10.76-0.98 (0.83), II 1.62-1.88 (1.75), III 1.20-1.37 (1.32), IV 1.02- 1.10 (1.06). Rostrum total 1. 2.22-2.40 (2.29), IV 0.54- 0.60 (0.56). Pronotum a.w. 1.00-1.10 (1.00), p.w. 1.30- 1.42 (1.36), 1. 0.72-0.84 (0.78). Wing pad 0.70-1.00 (0.86). Metafemur 1. 1.68-1.90 (1.82), w. 0.36-0.40 (0.39). Meta- tibia 1. 2.12-2.34 (2.25). Metatarsus 10.40-0.44 (0.42), II 0.52-0.58 (0.55). DAGO 0.115-0.132 (0.123). n=10 in all cases.

Material examined: 2 ex., Bucks, Fingest, 25.VII.59, GEW. 15 ex., Dumfs, Bodesbeck Farm, 20.VII.76, GMcG. 45 ex., Yorks, Malham Tarn, 21.VII.77, Juncus sp., Molinia caerulea, GMcG.

Remarks: This species is very common in grass upland habitats such as Malham Tarn and Bodesbeck Farm. The body of the Vth instar nymph is rather broader and squatter than that of the closely related S. laevigatum, which prefers lowland and more southerly habitats. - 429 - Stenodema laevigatum (L.) (S. Stenodema s.s.) Figs. 19:214,215; 39:3; 62

Green. Head and nota with longitudinal, lateral brown stripes. Antennae brown, IV with red tinge. Dorsal and ventral surfaces of abdomen with lateral red stripes, half way between body midline and margins. Legs light brown with faint spots. Claws fuscous. Head with faint longitudinal furrow posteriad. Pronotum with posterior margin indented medially. Dorsal body hairs short, black, spinous. Antennae strongly pubescent, segment I with hairs long, simple. DAGO double, weakly sclerotised with anterior bar sinuate. Claws striate. Pulvilli small. Parempodia flap-like. Trichobothria 7:6, trichomae present. Bothria recessed.

Measurements: Total body 1. 6.58-8.08 (7.35). Head W. 0.98-1.04 (0.99), 1. 1.02- 1.10 (1.06), vertex 0.46-0.56 (0.51). Antenna total 1. 5.42-6.48 (6.22), I 1.10-1.26 (1.18), II 2.32-2.48 (2.41), III 1.48-1.60 (1.54), IV 1.08- 1.24 (1.18). Rostrum total 1. 2.58-2.78 (2.67), IV 0.66- 0.70 (0.68). Pronotum a.w. 0.96-1.08 (1.00), p.w. 1.20- 1.34 (1.25), 1. 0.84- 0.90 (0.86). Wing pad 1.10-1.20 (1.16). Metafemur 1. 2.24-2.48 (2.35), w. 0.40-0.48 (0.43). Meta- tibia 1. 2.68-3.00 (2.83). Metatarsus 10.38-0.50 (0.46), II 0.48-0.60 (0.55). DAGO 0.110-0.125 (0.116). n=10 in all cases.

Material examined: 14 ex., Herts, Gt. Field, 12-17.VIII.53, Dactylis sp., TRES. 10 ex., London, Hampstead Heath, 25.VI.76, GMcG. 2 ex., Berks, Silwood Park, 16.VII.76, GMcG. 3 ex., Bucks, Denham, 27.VII.76, GMcG. 6 ex., London, Putney Heath, 4.VIII.76, GMcG.

Remarks: This species is common on low and southern grass- land, and is distinguished from S. calcaratum and S. trispin- osum by not having spines on the metafemora, and from S. holsaturn by being larger and more elongate. The nymphs feed on various grasses, Alopecurus pratensis, Phleum pratense and Festuca rubra (Southwood and Leston, 1959). Butler (1923) and Buczek (1959) have previously described the nymphs. ;; insert- Elongate, fusiform. - 430 - Notostira elonEata (Geoffroy) Figs. 8d; 19:216,217; 39:4; 62

Light green. Antennae ochreous. Head with central grey line merging into faint black furrow on vertex. Pronotum with two lateral brown stripes bordered with green, and a medial brown stripe with or without a pale central stripe. Meso- and meta-thoracic nota largely light brown to brownish- green. Abdomen with central green stripe and two lateral grey stripes. Legs light brown. Claws and tarsal apices black. Elongate, fusiform. Dorsal body hairs long and short, simple. DAGO double with anterior bar sinuate and with pigmented patches posteriad and anteriad. Claws striate. Pulvilli small, adpressed. Parempodia striate, flap-like. Trichobothria 9:8, trichs short, trichomae very weak. Bothria slightly recessed.

Measurements: Total body 1. 6.66-7.74 (7.34). Head w. 0.94-1.10 (1.03), 1. 1.06-1.20 (1.13), vertex 0.58-0.71 (0.64). Antenna total 1. 6.64-7.86 (7.04), I 1.30-1.46 (1.33), II 2.26-2.92 (2.45), III 1.78-2.10 (1.95), IV 1.26- 1.46 (1.32). Rostrum total 1. 1.98-2.24 (2.13), IV 0.38- 0.44 (0.40). Pronotum a.w. 0.92-1.12 (1.02), p.w. 1.20- 1.36 (1.27), 1. 0.79-0.96 (0.85). Wing pad 1.00-1.20 (1.10)n=9. Metafemur 1. 2.52-2.96 (2.64), w. 0.33-0.44 (0.40). Meta- tibia 1. 2.84-3.36 (2.95) . Metatarsus 10.50-0.60 (0.54), II 0.62-0.70 (0.66). DAGO 0.092-0.109 (0.104). n=10 in all cases except where shown.

Material examined: 2 ex., Herts, Gt. Field,25.VI.53, TRES. 6 ex., Herts, Gt. Field, 17.IX.53, TRES. 4 ex., Herts, Gt. Field, 26.IX.54, TRES.

Remarks: The nymphs of this bivoltine species give the appear- ance of having three longitudinal dark lines on the head, four lines on the pronotum, meso- and meta-thoracic nota, and three on the abdomen. This species shares a high trichoboth- rial complement with the genus Leptopterna. Butler (1923) described the last instar nymph as being similar to S. laevigatum but having the forehead produced beyond the clypeus, which is vertical. - 431 - Megaloceraea recticornis (Geoffroy) Figs. 19:218,219; 39:5; 62

Green. Nota with lateral brown bands. Tarsi, claws and apical half of rostral segment IV infuscate. Very slender, elongate. Antennae very long. Dorsal body hairs stout, blunt, short, black. Ventral body hairs pale, slender, simple. DAGO double, weak with anterior bar sinuate. Claws slender, slightly curved. Pulvilli small to medium, rugulose, adpressed. Parempodia flap-like, pointed. Trichobothria 7:6, trichomae weak. Bothria slightly recessed.

Measurements: Total body 1.-7.00-8.66 (7.67). Head w. 0.90-1.04 (1.00), 1. 1.00-1.12 (1.05), vertex 0.50-0.64 (0.57). Antenna total 1. 7.67-8.28 (7.95)n=8, I 1.16-1.36 (1.27), II 2.44-2.80 (2.63), III 2.88-3.12 (2.97), IV 1.08- 1.20 (1.12) n=8. Rostrum total 1. 2.50-2.76 (2.65), IV 0.62- 0.70 (0.65). Pronotum a.w. 0.96-1.04 (1.01), p.w. 1.20- 1.36 (1.28), 1. 0.70-0.84 (0.78). Wing pad 1.10-1.20 (1.16). Metafemur 1. 2.72-3.00 (2.88), w. 0.34-0.42 (0.37). Meta- tibia 1. 3.08-3.56 (3.35). Metatarsus I 0.38-0.50 (0.48), II 0.52-0.58 (0.56). DAGO 0.086-0.103 (0.099). n=10 in all cases except where shown.

Material examined: 11 ex., Herts, Harpenden, 29.VI.55, TRES. 1 ex., Sussex, Watlington, -.VII.67, GEW. 5 ex., Sussex, Camber Sands, 15.VII.77, GMcG.

Remarks: Butler (1923) mentioned the nymphs as being similar to those of Notostira elongata, but with longer antennae. Southwood and Leston (1959) said that the green lymphs were characterised in later instars by their long antennae and two brown bands on the thoracic margins. The antennae are the longest in the Stenodemini, (see fig. 63). This species was described new to the Eastern United States by Slater (1956) who gave measurements and descriptions for all instars. - 432 -

Trigonotylus ruficornis (Geoffroy) Figs. 19:220,221; 39:7; 62

Pale green. Antennal segment I with red tinges, II and III faintly red, IV with strong red tinge. Head with dark, longitudinal stripe, nota with four brown, longitudinal stripes. Abdomen with three orange-red longitudinal stripes. Small, slender, fusiform. Dorsal body hairs long, blunt (max 1. 0.05) and short, simple (max 1. 0.02). Ventral body hairs slender, pale, simple. DACO single or double, pale, slit-like, anterior bar curved or slightly sinuate. Claws slender, faintly striate. Pulvilli small. Parempodia flap- like, pointed, striate. Trichobothria 6:6, trichomae very weak. Bothria slightly recessed..

Measurements: Total body 1. 4.12-4.72 (4.42). Head w. 0.07- 0.82 (0.77), 1. 0.72-0.84 (0.78), vertex 0.40-0.44 (0.41). Antenna total 1. 3.72-4.46 (4.12), I 0.52-0.64 (0.58), II 1.28-1.60 (1.48), III 1.40-1.60 (1.50), IV 0.52-0.62 (0.57). Rostrum total 1. 1.22-1.56 (1.43), IV 0.30-0.40 (0.37). Pronotum a.w. 0.70-0.88 (0.79), p.w. 0.88-1.04 (0.93), 1. 0.44-0.56 (0.50). Wing pad 0.76-0.96 (0.84). Metafemur 1. 1.30-1.56 (1.44), w. 0.24-0.32 (0.29). Metatibia 1. 1.60- 1.90 (1.79). Metatarsus 10.26-0.32 (0.29), II 0.38-0.42 (0.40). DAGO 0.069-0.090 (0.082). n=10 in all cases.

Material examined: 11 ex., Herts, Harpenden, 29.VI.55, TRES. 2 ex., Berks, Silwood Park, 3.VII.77, GMcG.

Remarks: Buczek (1959) described the nymphs and Southwood and Leston (1959) gave Dechampsia flexuosa, Festuca rubra and Agrostis tenuis as the usual host plants. - 433 -

Teratocoris antennatus (Boheman) Figs. 19:222,223; 39:8; 62

Pale green with pink markings and light reddish-brown with faint green abdominal tinge. Antennal segment I, apex of segment II with red tinge. Head, nota, abdomen with thin,red, longitudinal, medial stripe. Abdomen with two lateral, longitudinal red stripes. Femoral apices with red tinges. Claws, apical half of tarsal segment II infuscate. Small, slender, subfusiform. Rostrum short reaching just past pro- thoracic coxae. Dorsal body hairs short, black, simple. Ventral body hairs long, slender, pale. DAGO single or weakly double, pale, anterior bar straight. Claws slender, faintly striate. Pulvilli small. Parempodia flap-like. Trichobothria 7:6, trichomae marked, compact. Bothria well recessed.

Measurements: Total body 1. 3.72-5.08 (4.29). Head w. 0.74- 0.84 (0.80), 1. 0.58-0.70 (0.64), vertex 0.36-0.44 (0.39). Antennal total 1. 4.38 n=1, 10.72-0.80 (0.78), II 1.18- 1.38 (1.27), III 1.18-1.20 (1.19) n=3, IV 1.00 n=1. Rostrum total 1. 1.13-1.28 (1.18), IV 0.30-0.36 (0.34) n=5. Pro- notum a.w. 0.68-0.78 (0.72), p.w. 0.80-1.01 (0.89), 1. 0.51-0.62 (0.56) . Wing pad 0.50-0.68 (0.53) n=3, 6 0.40- 0.42 (0.41) n=3. Metafemur 1. 1.24-1.50 (1.41), w. 0.24-0.27 (0.26). Metatibia 1. 1.34-1.62 (1.52). Metatarsus I 0.24-0.28 (0.25), II 0.36-0.42 (0.40) . DAGO 0.074-0.086 (0.079) n=6 except where shown.

Material examined: 4 ex., Mddsx, Hounslow, -.VI.53, GEW. 2 ex., Mddsx, Hounslow, -.VI.54, GEW.

Remarks: Butler (1923) and Kullenberg (1944) have described the nymphs and Southwood and Leston (1959) gave Scirpus maritimus as the main host plant and also Juncus gerardii Glyceria fluitans and Schoenoplectus tabernaemontani. The shape of the first anten- nal segment with slight constriction subapically is charac- teristic of the genus Teratocoris. The d nymphs examined had shorter wing pads than those of the ~~. - 434 - Teratocoris saundersi Douglas and Scott Figs. 18e,f; 19:228,229; 40:1; 62

Entirely yellowish-green. Antennae with red tinge except at base of segment I. Eyes red. Apex of femora with reddish tinge. Small, slender, subfusiform. Dorsal body hairs short, black, spinous. Ventral body hairs shorter, simple. DAGO single or weakly double, slit-like, anterior bar straight. Claws slender striate. Pulvilli small. Parempodia flap-like. Trichobothria 8:7, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.40-4.76 (4.61). Head w. 0.80- 0.90 (0.85), 1. 0.62-0.68 (0.65), vertex 0.43-0.50 (0.47) . Antenna total 1. 4.92-5.19 (5.11), I 0.88-0.96 (0.90), II 1.76-1.86 (1.82), III 1.22-1.36 (1.29), IV 1.06-1.11 (1.09). Rostrum total 1. 1.24-1.26 (1.25), IV 0.34-0.36 (0.35). Pronotum a.w. 0.74-0.80 (0.78), p.w. 0.96-1.00 (0.97), 1. 0.56 -0.60 (0.58) . Wing pad 0.52-0.60 (0.55) . Metafemur 1. 1.76 -1.90 (1.81), w. 0.29-0.32 (0.30). Metatibia 1. 1.90- 2.00 (1.93). Metatarsus I 0.34-0.36 (0.35), II 0.48-0.52 (0.50). DAGO 0.078-0.082 (0.080). n=4 in all cases.

Material examined: 5 ex., Dumfs, Bodesbeck Farm,. 10.VII.78, Carex and Juncus spp., GMcG.

Remarks: The nymphs of the genus Teratocoris are all very similar. - 435 -

Teratocoris viridis Douglas and Scott Figs. 19:224,225; 40:2; 62

Green or bright green. Antennal segment II, III, IV and apex of segment I with red tinge. Eyes red. Claws, apical half of tarsi and rostral apex black. Small, slender, sub- fusiform. Dorsal body hairs short, black, spinous, becoming slender on posterior abdominal segments. Ventral body hairs long, slender, simple. DAGO single or weakly double, anterior bar straight. Claws slender, slightly curved, faintly striate. Pulvilli small. Parempodia flap-like, striate. Trichobothria 7:6, trichomae marked, compact. Bothria recessed.

Measurements: Total body 1. 4.36-4.74 (4.50). Head w. 0.83- 0.92 (0.88), 1. 0.68-0.73 (0.71), vertex 0.46-0.48 (0.47). Antenna total 1. 4.12 -4.44 (4.24), I 0.74-0.88 (0.79), II 1.42-1.54 (1.45), III 1.03-1.10 (1.07), IV 0.89-0.96 (0.93). Rostrum total 1. 1.23-1.35 (1.29), IV 0.34-0.38 (0.35). Pronotum a.w. 0.78-0.86 (0.82), p.w. 0.96-1.09 (1.03), 1. 0.58-0.66 (0.62) . Wing pad 0.60-0.82 (0.65) . Metafemur 1. 1.60-1.83 (1.69), w. 0.32-0.34 (0.33). Metatibia 1. 1.66- 1.82 (1.74). Metatarsus I 0.32-0.36 (0.33), II 0.44-0.48 (0.46). DAGO 0.086-0.092 (0.089). n=4 in all cases.

Material examined: 4 ex., Outer Hebrides, South Uist, Loch , 28.VI.77, ARW.

Remarks: Southwood and Leston (1959) gave Carex spp. as the probable host plants for this species. - 436 - Teratocoris caricis Kirkaldy,1909 = elegans Woodroffe, 1967. (Woodroffe, 1967 & 1969) Figs. 19;226,227; 62

Bright green. Antennae with red tinges, segment I green basally with red tinge apically. Head with central, pale, fuscous stripe. Pronotum green with pale fuscous stripes down lateral margins. Wing pads green with pale fuscous markings. Femora green with red tinges apically. Tarsi fuscous with apex of segment II and claw becoming black. Coxae with faint fuscous bands. Small, slender, subfusiform. Antennae long, slender. Rostrum reaching nearly to mesothora- cic coxae. Legs slender. Dorsal body hairs very sparse, black, very short, spinous, becoming less sparse on femora, tibiae and antennae. Ventral body hairs pale, simple. DAGO single or weakly double, broad, slit-like, anterior bar straight. Claws slender,striate. Pulvilli small. Parempodia flap-like. Trichobothria 9:7, trichomae present. Bothria recessed.

Measurements: Total body 1. 4.44-4.80 (4.64). Head w. 0.84- 0.94 (0.87), 1. 0.62-0.70 (0.66), vertex 0.46-0.56 (0.50). Antenna total 1. 4.76-5.18 (5.00) n=4, I 0.84-0.92 (0.89), II 1.66-1.90 (1.79), III 1.16-1.30 (1.22), IV 1.04-1.08 (1.06) n=4. Rostrum total 1. 1.26-1.31 (1.27), IV 0.34-0.39 (0.36). Pronotum a.w. 0.76-0.87 (0.79), p.w. 0.98-1.07 (1.02), 1. 0.58-0.64 (0.60). Wing pad ? 0.54-0.68 (0.61) n=3, 0.76-0.82 (0.78) n=5. Metafemur 1. 1.66-1.94 (1.80), w. 0.29- 0.37 (0.32). Metatibia 1. 1.74-2.02 (1.90). Metatarsus I 0.32-0.38 (0.35), II 0.47-0.52 (0.49). DAGO 0.01 (0.01) n=4. n=8 except where shown.

Material examined: 13 ex., Perths, Rannoch School, 24.VI.78, Juncus sp., GMcG.

Remarks: The adults and nymphs of this species are found in freshwater marshland and are best captured by searching among the bases of clumps of rush and sedge. Three macropterous d adults and two brachypterous adults were found, the Vth instar ? nymph having correspondingly short wing pads not reaching to the DAGO, the d with long wing pads reaching to or a little beyond the DAGO. - 437 -

Leptopterna dolabrata (L.) Figs. 8e; 19:230,231; 40:3; 43b; 62

Green or greenish-yellow in young instar nymphs or olive- grey in later instar nymphs with dark brown or black markings. Antennal segment I and basal two-thirds of segment II brownish-red, remainder brownish-black. Head with central dark markings and thin white ecdysial line. Eyes dark red with dark markings posteriad. Rostral segments III and IV infuscate. Pronotum pale with calli and posterior margin dark brown and with thin white ecdysial line. Wing pads pale laterally and medially, otherwise dark brown with anterior and posterior margins strongly brownish-black. Abdomen green with brownish-grey patches on each segment, one on either side of thin, white, medial stripe. Femora pale, tibiae brown to brownish-black. Tarsi and claws infuscate. Dorsal body hairs strong, black, simple (max.l. 0.15). Ventral body hairs pale, slender, simple (max.l. 0.16). DAGO double, broad, dark, anterior bar sinuate and with pig- mented patches anteriad and posteriad. Claws thick basally, striate. Pulvilli small, rugulose. Parempodia flap-like, pointed, striate. Trichobothria 9:8, trichomae present. Bothria slightly recessed.

Measurements: Total body 1. 6.16-7.58 (7.83). Head w. 1.12- 1.24 (1.18), 1. 0.98-1.18 (1.08), vertex 0.56-0.72 (0.64). Antenna total 1. 6.12-6.52 (6.36) n=9, I 0.96-1.06 (1.01), II 2.40-2.72 (2.58), III 1.64-1.80 (1.71), IV 1.00-1.08 (1.03) n=9. Rostrum total 1. 2.38-2.76 (2.55) n=8, IV 0.60-0.66 (0.62). Pronotum a.w. 1.10-1.28 (1.16), p.w. 1.28-1.58 (1.47), 1. 0.80-0.84 (0.82). Wing pad ? 0.32-0.64 (0.48)n=6, 1.04-1.22 (1.13)n=4. Metafemur 1. 2.08-2.32 (2.21), w- 0.36-0.50 (0.44). Metatibia 1. 2.68-2.90 (2.81). Metatarsus I 0.44-0.50 (0.47), II 0.58-0.66(0.62). DAGO 0.11-0.15 (0.13). n=10 except where shown.

Material examined: 12 ex., Herts, Harpenden, 10-17.VI.53, Dactylis glomerata, TRES. 18 ex., Berks, Silwood Park, 17.VI.77, GMcG.

Remarks: Southwood and Leston (1959) gave information on - 438 -

the biology, distribution and host plant preferences, including Phleum pratense, Agropyron repens, Alopecurus pratensis, Dactylis glomprata and Holcus lanatus. Butler (1923) and Buczek (1959) described the nymphs. The wing pads of the Vth instar d nymphs are twice as long as those of the nymphs. The effect of this species on Kentucky blue grass (Jewett & Townsend,1947) and on the yield and quality of wheat (Rautapaa, 1970) has been investigated. The energetics and population dynamics of this species have been studied McNeill (1971 & 1973) . - 439 -

Leptopterna ferrugata (Fallen) Figs. 19:232,233; 40:4; 62

Pinkish-pale brown with brown markings and medial white stripes. Antennal segment I, basal two-thirds of segment II pale, remainder dark with red tinge. Head with central brown large 'V' marking. Eyes dark with dark brown markings posteriad. Pronotum pale with calli dark brown. Wing pads with lateral margins pale, central dark brown stripe on each pad extending laterally to the internal margins. Abdomen with brown markings giving appearance of broad, brown band running length of abdomen, extending laterally to connexivum and broken by central white line and row of patches on either side. Legs light brown. Femora with dark spots. Tarsi, claws, tibial apices and rostral segments III and IV infuscate. Dorsal body hairs stout, black, simple (max.l. 0.13). Ventral body hairs pale, simple (max.1. 0.14). Femora, tibiae with long spines. DAGO double, sclerotised, anterior bar sinuate. Claws elongate, striate. Pulvilli small to medium, adpressed. Parempodia flap-like, pointed, striate. Trichobothria 9:8, trichomae present. Bothria slightly recessed.

Measurements: Total body 1. 5.83-7.16 (6.78). Head w. 0.90- 1.14 (1.06), 1. 0.92-1.16 (1.03), vertex 0.48-0.64 (0.56). Antenna total 1. 4.95-5.54 (5.28) n=9, I 0.84-1.00 (0.93), II 1.83-2.20 (2.05), III 1.40-1.66 (1.49), IV 0.80-0.90 (0.83) n=9. Rostrum total 1. 2.04-2.65 (2.34), IV 0.52-0.70 (0.60). Pronotum a.w. 0.94-1.22 (1.08), p.w. 1.16-1.52 (1.41), 1. 0.64-0.90 (0.77). Wing pad 0.34-0.60 (0.42) n=6,d 0.74- 1.10 (0.99)n=4. Metafemur 1. 1.70-2.10 (1.90), w. 0.34-0.49 (0.43). Metatibia 1. 2.08-2.56 (2.30). Metatarsus I 0.38-0.50 (0.44), II 0.50-0.60 (0.54). DAGO 0.080-0.109 (0.096). n=10 except where shown.

Material examined: 11 ex., Herts, Harpenden, 29.VI.55, TRES. 7 ex., Berks, Silwood Park, 17.VI.77, GMcG.

Remarks: This species is very similar to L. dolabrata, but is found in drier conditions, has less dark colouration on the pronotum, shorter antennae and is more slender. The - 440 -

DAGO does not have such strong pigmentation. Butler (1923) described the nymphs briefly and Southwood and Leston (1959) gave the host plants as Festuca rubra, Agrostis tenuis, Deschampsia flexuosa and Poa pratensis. As in L. dolabrata there is sexual dimorphism in the nymphal wing pads. - 441 -

DISCUSSION

It is difficult to examine broad phylogenetic implications when dealing with the restricted fauna of a small geographic region as in the present study. However, valuable informat- ion indicating relationships not supported by adult charac- ters can be obtained by examination of nymphal characters.

The phylogeny of adult Miridae has been examined previously by various authors (Reuter, 1910; Knight,1923; Slater, 1950; Carvalho, 1952; Wagner, 1955, Kelton, 1959; Leston, 1961 and Schuh, 1974). Many of the published phylogenetic schemes have been based on only a part of the character spectrum due to lack of available information. However, Cobben (1978) has provided an excellent evolutionary scheme for the Heteroptera, and Schuh (1974) gave an extensive phylogenetic analysis of the world ant-mimetic tribes of the Orthotylinae and Phylinae. Schuh's work dealt only with adults, but Cobben (1978) has shown that nymphal characters are valuable in contructing evolutionary schemes and has such nymphal characters as numbers of omuiiatidia, tarsal seg- ments, dorsal abdominal glands, as well as the condition of the first abdominal spiracle. In the present study the nymphs of British Miridae have been examined in detail, using trichobothria, gland openings and pretarsal structures. In addition one tribe, the Stenodemini, has been examined using numerical taxonomic techniques. The latter proved very powerful and were able to distinguish all species on the basis of the morphological characters used. The Stenodemini have evolved to share a grass-feeding habit and have acquired an elongate, fusiform body shape. Numerical techniques proved able to classify adequately this seemingly homogeneous tribe, and their use in other, more hetero- geneous, taxa should be even more rewarding. Enough morpho- metric data has been collected to allow this type of study to be extended to most of the other tribes.

Heteropteran pretarsal structures have been reviewed by Cobben (1978) and the mirid pretarsus was studied by Schuh (1976), using adult material. The present study on nymphs has shown that the pretarsus does not change in-basic - 442 - structure throughout ontogeny, although Cobben(1978) re- corded drastic ontogenetic change of the pretarsus in (). It would appear that, in general, the mirid nymphal pretarsus does not offer any additional taxonomic characters to those of the adult. Ontogenetic change in pretarsal structures might be expect- ed to occur in taxa where the larval and adult life styles differ markedly. However, as the nymphal life style is not significantly different from that of the adults in mirids, it is not surprising that little ontogenetic change has been observed.

There appears to be confusion about the pretarsal structure of Bryocorinae and several authors have commented on the need for further work on this subfamily. In the present study Monalocoris filicis and Bryocoris pteridis were exam- ined (fig. 5a). The Monalocoris studied differed from the one illustrated by DeCoursey (1971). The con- fusion may have arisen from the inability of optical micro- scopy to determine accurately the origin and thus the nature of the structures. The resolving power of the scanning electron microscope now enables a more accurate appraisal of the pretarsus. The poorly defined bryocorine taxonomy adds to the confusion. Akingbohungbe (1974) stated that the Bryocorinae have large flap-like pulvilli, arising from the ventral surface of the claw. Close examination of the British taxa shows these structures to arise from or near the unguitractor plate. Thus they cannot be termed pulvilli, but should rather be termed parempodia (terminology of Schuh,1976). They appear similar to the flap-like parem- podia seen in Orthotylini and Pilophorini. Schuh (1976) did not consider that they were parempodial in nature because they are not set in a socket -.a characteristic of mechano- sensory setae from which p'arempodia are supposedly derived. He concluded that they are pseudopulvilli and distinct from true pulvilli in that they arise not from the claws them- selves, but rather laterad of the parempodia on the ungui- tractor plate. Cobben (1978), in agreement with Schuh (1976), stated that'the paired bladder-like pretarsal outgrowths'were neither true pulvilli or parempodia, and further figured vestigial parempodia for Bryocoris pteridis. These latter - 443 - were not seen by the present author in Bryocoris pteridis or Monalocoris filicis. In this study it is considered that the large flap-like structures arising from the apical portion of the unguitractor plate are parempodia and not pseudopulvilli. However, confirmation of the real nature of these structures awaits histological examination of the pretarsus to determine whether or not they are sensory.

It is apparent that the Bryocorinae present a confused picture. Cobben (1968) stated that the pest genus Helopeltis, currently placed in the Bryocorinae, has egg characters quite different from the 'typical' Bryocorinae and its pretarsus is not like that of Bryocoris or Monalocoris. Schuh (1976) has pointed out that the Bryocorinae have been neglected because of an almost totally tropical distribution, which has led to an assessment of relationships based on the non-representative fauna of the temperate regions of the northern hemisphere. The sub- family requires careful study and evaluationcf more charac- ters before a proper assessment of its relationships can be made.

In 1976, Schuh proposed major alteration to the ci assification of the Miridae. The Orthotylinae of authors was recognised as a tribe within the Phylinae. The Deraeocorinae of authors was recognised as a tribe within the Mirinae and the Bryocorinae was redefined to include the Dicyphinae of authors.

The Dicyphinae was previously placed in the subfamily Phylinae by Carvalho(1952), but given subfamily status by Leston (1961) and others (see Schuh, 1976). Schuh has related the Dicyphinae to the Monaloniina and Bryocorinae on the basis of synapomorphies in the pseudopulvilli and trichobothrial number and pattern, and Cobben (1968) further related them to the Monaloniina by the presence ofiespiratory horns on the eggs. The nymphs examined in the present study support the elevation of the group to subfamily level.

Schuh (1976) considered the Bryocorini and Dicyphini to possess pseudopulvilli (as already discussed for Bryocorinae). - 444 -

The structure of the dicyphine claw appears very different from that of Bryocoris or Monalocoris. The claw structures can be seen in Figures 5a, 6a-f and 55-56. The claw is elongate with a basal projection which bears a large flap- like pulvillus. The setiform (hair-like) parempodia (often striate) can be clearly seen arising from the unguitractor plate. It should be noted here that Campyloneura virgula, at present included in the Dicyphinae, is unusual in having extensive polygonal surface sculpturing and very long tarsal guard setae (fig. 56), characters which point to a relationship with the Bryocorinae. Campyloneura is unusual in other respects, being parthenogenetic and having pretarsi subapically positioned.

The subfamily rank of the Dicyphinae is supported further by other nymphal features. The adults of Bryocorinae, Dicyphinae, Mirinae and Deraeocorinae possess a rounded pronotal collar - this is usually absent in the Orthotylinae and totally absent in the Phylinae. The nymphs of Dicyphinae are unique in possessing a pronotal collar (although to a lesser degree of development than the adults). The long second tarsal segment, the expanded abdominal spiracular openings (this latter character shared with the nymphs of Halticocorini) and the structure of the DAGO and gland sac also indicate that the Dicyphinae are not very closely related to other subfamilies. The DAGO is very small with a single opening and a small gland sac. (figs. 41a, c; 42f; 43e; 27; 28). The exception again is C. virgula, whose DAGO appears double, supporting its affinity with another taxon. The trichobothrial type and pattern in the Dicyphinae (pp.63-64) are unique in the nymphs examined and although with a superficially similar bothrial type to the Bryocorinae, the lack of fused trichomae in•the latter taxon and its much reduced trichobothrial number denies any real relationship.

The dicyphine pulvillus has a completely different orientat- ion and probable origin to that in the Phylinae ( a relation- ship suggested by Carvalho, 1952). The phyline genus Macrotylus has two species in Britain, paykulli and solitarius, which have pretarsal structures differing from those of all other phylines examined, and bearing a strong resemblance to - 445 - those in the Dicyphinae (fig. 49). The pulvilli are large, free along their entire length and originating from a basal projection (although not as large as in the Dicyphinae) of the claw. Seidenstlicker (1967) has already pointed out that Macrotylus contains some species with phyline-type claws and some with dicyphine-type claws. It is considered that the pulvilli arising from the basal projection of the claw in the Dicyphinae and MacroLylus is in no way related to the pseudopulvillus (of Schuh, 1976) in the Bryocorinae - rather they are unique structures, derived from pulvilli which are normally situated on the ventral claw surface, in response to substrate or other factors. The possession of similar looking strucutres in Macrotylus and the dicyphines may reflect an environmental adaptation rather than any close phylogenetic relationship. The function of the pulvilli and pseudopulvilli (of Schuh) is not known, but has been discussed by Wagner (1955), Seidenstiicker (1967) and Schuh (1976). The hypothesis that species with elongate, free pulvilli, as inthe Dicyphinae, live on plants with sticky hairs, the claws structures representing an adaptation to the substrate, has been examined by these authors. Southwood (1972) mentioned that some species use trapped insects on such plants as small, but important, secondary food sources. It is possible that some species avoid being trapped by sticky secretions or glandular plant hairs by behavioural adaptations, i.e. slow movement and modified preening (Dolling, personal communication).

Schuh (1976) pointed out that there are taxa that possess similar'pulvilli' (Schuh's inverted commas), but do not live on sticky plants. In considering the function of pulvilli, he also stated that the form of the pulvilli in Eminoculus drosanthemum Schuh (Phylinae) was suggestive of some adhesive function, as they covered the entire ventral surface of the claw. Such inflated pulvilli occur in phyline taxa examined in the present study (figs. 5d; 53), being most clearly seen in Lopus decolor, where the pulvilli are very large and would seem to prevent any part of the claws contacting the substrate in normal use. Such structures have not been found in other subfamilies and further SEM and ultrastructural study will be valuable. - 446 -

Following the work of Kelton (1959) on d genitalia and Slater (1950) on genitalia, Schuh (1976) placed the Deraeocorinae in the Mirinae as a tribe on the basis of the adult pretarsus. Leston (1961) also proposed a link between the Deraeocorinae and the Mirinae following the work of Kelton and Slater, together with evidence from his work on chromosomes, testis follicle numbers and wing ven- ation. In the present work it is considered that the nymphal pretarsus is not sufficiently similar or dissimilar to that of the Mirinae to allow discussion. However, the parempodia of the deraeocorines examined are strap-like and and not flap-like as in the Mirinae, and the pulvilli are much altered and not similar to the lobe-like pulvilli on the ventral surface of the mirine claw. The trichomae of the Deraeocorinae are moreover diffuse rather than compact as in the Mirinae. When studying the DAGOs of the two taxa there appears to be a similarity in that both groups possess double openings, but this is much more marked in the Deraeocorinae.

Reuter (1910) and Knight (123) considered the Phylinae to be among the most primitive Miridae, but Schuh (1974) considered them to be among the most highly evolved, the 6 genitalia being structurally distinct from those of all other Miridae and having a unique and complex functional relationship. The dorsal hair types in the nymphs of Miridae , may provide further insight into the relationships of the group. Ignoring the taxa with very specialised and unique hair types such as Harpocera thoracica (Phylinae) (fig. 1f), Capsus ater (Mirinae) (fig. 3a) and the Hallodapini (Phylinae) (fig. 2d), the Phylinae appear to possess a derived nymphal hair type. This hair type is usually short, trifurcate or apically multiply branched (coronate) and occurs over the entire dorsal surface. The plesiomorphous form is here considered to be simple (setiform) hairs without any apical branching. The other tribes examined possess predominantly simple hairs or a low proportion of bifurcate or trifurcate hairs, restricted to certain parts of the dorsal surface. A study of the nymphal dorsal hair types in other families of Heteroptera may yield valuable information on inter- family relationships. - 447 -

Schuh (1974) stated that the Hallodapini have generally included all Phylinae that are ant-mimetic, and he con- sidered them to form a monophyletic unit within the sub- family, basing this view on the adult characters of ant- mimetic facies, flattened pronotal collar, hemelytral maculae and the structure of the ð genitalia. The unique hair type,seen in the nymphs of the three British taxa Hallodapus rufescens, H. montandoni and Systellonotus triguttatus, namely swollen-ended, elongate setae, support this. Schuh also recorded this group as .having the first stridulatory mechanism described in the Miridae. The lateral corial margin is described as bearing a 'stridulitrum' of fine teeth, the plectrum occurring on the inner surface of the metafemur. Trichophthalmocapsus, Laemocoris and some species of Hallodapus (including those found in Britain) are described as having these structures, but convincing evidence of stridulation was not given. It is interesting that the nymphs of H. rufescens and H. montandoni appear to possess the metafemoral structure (fig. 2e) which extends to the lateral abdominal cutcle in the area of the metafemur (an area which cannot contact the corial margin in the adult). The'stridulitrum' of Schuh does not appear to be possessed by the British Hallodapus adults, although the corial margin is much thickened. The structure is certainly very unusual and is not possessed by Systellonotus triguttatus, although it shares a similar ground living habit to the species studied by Schuh. The possibility of stridulation is not, however, ruled out.

Schuh (1976) stated that, apart from the lamelliform (flap- like) parempodia, no really strong apomorphic characters have been found that hold the Orthotylini together as a group. It is proposed here that the structure of the ungui- tractor plate, together with possession of flap-like parem- podia, may be suitable. All Orthotylines examined had broad unguitractor plates with two lateral, curved rows of scales, the medial row,apparently always present in mirines, being absent or reduced.

Future work,suggested by this study,includes examination of the trichobothria using transmission electron microscopy, - 448 -

together with behavioural and physiological studies, in order to determine their function in the Miridae and perhaps shed light on the various trichobothrial types found. A study of the nymphal secretions (where present) of the dorsal abdominal gland, using gas chromatography and other analyti- cal techniques, would be of value in suggesting relationships within the family and at higher taxonomic levels. The Bryocorinae requires an extensive analysis on a world-wide basis to clear up the taxonomic and phylogenetic confusion that exists about the group. Work on a broader scale should include study of the tracheal system of heteropteran families and spiracular structure as this may prove useful in delimiting relationships. - 449 -

Appendix A - List of collectors

The following initials appear in the descriptions:

EAB E.A. Butler* WRD W.R. Dolling DOD D. O'Donnell DMG Dr D.M. Glen EWG E.W. Grove s GMcG G.C. McGavin RH R. Harrington DL Dr.D. Leston MMM Dr M.M. Manson JHM J.H. Martin AMM Dr A.M. Massee* CWM C.W. Moffat MGM Dr M.G. Morris JGO J.G. Ottoson RGP Dr R.G. Paul ECP-C E.C. Pelham-Clinton TRES Professor T.R.E. Southwood RW Dr R. Waterston GWW G.W. Watson GEW G.E. Woodrof f e

deceased - 450 - Appendix B - Collecting sites

The collecting sites shown on the map were visited during 1975-1978, some of them several times.

1 Harpenden 2 Slough 3 Windsor Great Park 4 Chobham Conunon/Silwood Park/Longcross/Virginia Water 5 Richmond Park 6 Wimbledon Common/Putney Heath 7 Hampton Court 8 Hampstead Heath 9 Epping Forest 10 Denham 11 Abinger Hammer/Ranmoor Common 12 Boxhill 13 Rye/Camber Sands 14 Witley Wood (New Forest) 15 Milton Abbas 16 Bristol (Long Ashton) 17 Malham Tarn 18 Bodesbeck Farm 19 St Mary's Loch/Loch of the Lowes 20 Wanlockhdad/Leadhills 21 Edinburgh 22 Rannoch School 23 Kinloch Rannoch - 451 -

Figure 79 - Collecting sites

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9 ;.~'- ~~-9 N2-~~--- .

8 N1 ------~-

7 ------K H: I

H IL E 5: I 9 lOa

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8 or=l======t====l

o

4 Appendix C - Statistical data for the Stenodemini (Vth instar) Acetropis gimmerthali 1 (rostrum I) $.64 0.60 8.62 6.60 6.56 0.64 L58 0.48 6.60 0.52 (rostrum II) 0.58 0.48 0.44 0.58 0.40 0.48 0.44 0.44 0.58 0.48 (rostrum III) 0.56 0.58 0.58 0.54 0.46 0.52 0.52 0.44 0.46 0.50 4 (rostrum IV) 0.52 0.54 0.52 0.50-0.46 0.50 0.48 0.44 0.50.0.50 5 (total rostrum) 2.30 2.20 2.16 2.22 1.88 2.14 2.02 1.80 2.14 2.00 3 (antenna I) 0.64 0.66 0.66 0.64 0.60 0.64 0.64 0.58 0.68 0.60 7 (antenna II) 1.52 1.58 1.44 1.50 1.36 1.62 1.48 1.40 1.56 1.50 ! ,_ (antenna III) 0.72 0.70 0.70 0.72 0.76 0.74 0.70 0.78 0.72 0.72 ,„ 9 (antenna IV) 0.52 0.50 0.50 0.50 0.50 0.50 0.50 0.50 0.52 0.52 u, 10 (total antenna) 3.40 3.44 3.30 3.36 3.22 3.50 3.32 3.26 3.48 3.34 11(head width) 0.92 0.92 0.90 0.90 0.80 0.90 0.92 0.80 0.94 0.90 i 12 (vertex width) 0.64 0.64 0.64 0.64 0.60 0.62 0.66 0.52- 0.66 0.60 1 (Read length) 0.82 0.80 0.76 0.80 0.66 0.78 0.80 0.64 0.78 0.80 14(total body length) 5.83 5.50 5.41 5.25 5.00 5.58 5.25 5.08 5.58 5.25 15(metatibial length) 1.80 1.84 1.72 1.74 1.64 1.88 1.76 1.62 1.72 1.74 16(metafemoral length) 1.30 1.36 1.32 1.26 1.20 1.30 1.20 1.12 1.30 1.24 17(metafemoral width) 0.36 0.32 0.32 0.34 0.30 0.34 0.32 0.32 0.36 0.34 18(metatars. seg. I) 0.28 0.28 0.28 0.28 0.26 0.27 0.28 0.24 0.28 0.27 19(metatars. seg. II) 0.46 0.46 0.44 0.46 0.44 0:45 0.44 0.42 0.46 0.44 20(DAGO) .115 .115 .109 .103 .092 .115 .097 .103 .115 .115 21(pronotum ant. width) 1.00 1.00 0.96 1.00 0.80 0.96 0.96 0.84 1.00 1.00 22(pronotum post. width) 1.40 1.42 1.38 1.38 1.20 1.38 1.36 1.20 1.42 1.40 23(pronotum length) 0.64 0.64 0.64 0.62 0.58 0.64 0.58 0.50 0.62 0.60 24(metathor. wing pad lgth) 0.76 0.79 0.74 0.81 0.82 0.80 0.76 0.74 0.78 0.80 Appendix C - (continued)

Stenodema calcarattml S. trispinosum 1 L44 L50 8.52 0.54 0.50 8.62 8.60 8.64 L62 0.38 0.38 0.44 L44 0.44 0.44 8.46 8.48 8.50 0.48 2 0.52 0.50 0.46 0.46 0.50 0.52 0.54 0.46 0.46 0.44 0.46 0.50 0.46 0.44 0.48 0.50 0.50 0.48 0.52 3 0.52 0.46 0.54 0.54 0.48 0.48 0.54 0.50 0.48 0.54 0.40 0.36 0.40 0.44 0.52 0.42 0.46 0.44 0.40 0.44 4 0.50 0.50 0.48 0.46 0.50 0.58 0.48 0.54 0.48 0.48 0.42 0.46 0.46 0.48 0.40 0.44 0.48 0.46 0.44 0.46 5 1.92 2.02 1.98 1.98 2.00 2.16 2.18 2.06 2.10 1.64 1.66 1.80 1.82 1.82 1.78 1.90 1.88 1 82 1.90 6 0.92 0.86 0.86 0.80 0.84 0.80 0.84 0.88 0.76 0.82 0.78 0.76 0.80 0.76 0.89 0.80 0.78 0.80 0.78 0.74 7 1.96 1.86 1.82 1.80 1.96 1 72 1.78 1.88 1.62 0.84 1.94 1.76 1.70 1.70 1.84 1.72 1.80 1.76 1.70 1.66 8 1.08 1.12 1.06 1.04 1.08 0.94 1.08 1.14 0.94 1.10 1.04 0.96 1.00 0.94 1.06 0.98 1.02 0.98 0.96 0.86 9 0.72 0.76 0.76 0.74 0.74 0.68 0.74 0.80 0.72 0.76 0.70 0.70 0.70 0.70 0.76 0.78 0.74 0.72 0.64 10 4.68 4.60 4.50 4.38 4.62 4.14 4.44 4.70 4.04 4.52 4.46 4.18 4.20 4.10 4.46 4.28 4.34 4.16 3.88 11 0.92 0.96 0.90 0.88 0.88 0.82 0.92 0.90 0.82 0.90 0.92 0.90 0.92 0.92 0.96 0.90 0.94 0.94 0.96 0.94 in 12 0.42 0.58 0.50 0.52 0.50 0.48 0.52 0.54 0.50 0.54 0.50 0.52 0.50 0.52 0.54 0.52 0.54 0.52 0.54 0.52 13 0.92 0.96 0.92 0.88 0.90 0.84 0.90 0.92 0.86 0.88 0.90 0.88 0.90 0.90 0.94 0.94 0.90 0.92 0.90 0.90 ' 14 5.49 6.41 6.08 5.58 5.66 5.00 5.50 5.60 4.66 5.91 5.33 5.41 5.75 6.00 6.16 5.66 6.41 5.66 5.41 5.41 15 2.10 2.16 2.00 1.96 2.02 1.84 2.00 1.98 1.78 2.02 2.04 1.92 1.96 1.92 2.10 2.06 2.02 2.02 1.96 1.90 16 1.78 1.88 1.76 1.66 1.70 1.56 1.74 1.64 1.50 1.70 1.74 1.60 1.78 1.68 1.8C 1.74 1.80 1.74 1.74 1.64 17 0.36 0.34 0.34 0.34 0.32 0.30 0.34 0.32 0.32 0.33 0.40 0.38 0.40 0.38 0.42 0.40 0.40 0.40 0.42 0.40 18 0.38 0.38 0.34 0.34 0.36 0.34 0.36 0.34 0.30 0.34 0.36 0.34 0.34 0.32 0.36 0.36 0.34 0.34 0.32 0.34 19 0.52 0.52 0.48 0.48 0.52 0.50 0.50 0.52 0.46 0.50 0.54 0.52 0.50 0.52 0.54 0.52 0.54 0.52 0.52 0.52 20 0.11 0.12 0.10 0.10 0.10 0.09 0.12 0.09 0.10 .100 .100 .1.00 .115 .120 .100 .100 .100 .100 .100 21 0.90 0.96 0.90 0.88 0.90 0.86 0.94 0.92 0.88 0.96 0.96 0.94 0.94 0.96 1.00 1.00 1.02 0.98 1.04 1.00 22 1.10 1.36 1.12 1.16 1.14 1.04 1.18 1.14 1.10 1.20 1.22 1.24 1.32 1.30 1.36 1.24 1.30 1.30 1.36 1.28 23 0.76 0.76 0.70 0.70 0.70 0.62 0.72 0.70 0.66 0.70 0.66 0.72 0.74 0.74 0.74 0.62 0.76 0.72 0.72 0.72 24 1.24 1.20 1.08 1.10 1.08 1.00 1.12 1.06 1.04 1.08 1.04 1.06 1.12 1.08 1.14 1.14 1.08 1.14 1.06 1.02

Appendix C - (continued)

S. holsatum S. laevigatum 1 0.70 L60 L.58 0.60 0.60 0.60 0.58 0.62 0.62 0.60 0.60 0.60 0.64 0.66 0.70 0.70 0.70 0.72 0.62 0.58 2 0.58 0.60 0.56 0.56 0.58 0.56 0.60 0.52 0.54 0.58 0.72 0.74 0.70 0.74 0.72 0.72 0.72 0.72 0.52 0.72 3 0.56 0.54 0.56 0.54 0.54 0.54 0.54 0.54 0.52 0.52 0.64 0.66 0.62 0.58 0.62 0.66 0.64 0.60 0.82 0.60 4 0.56 0.60 0.58 0.58 0.58 0.54 0.54 0.56 0.54 0.56 0.66 0.68 0.68 0.68 0.68 0.70 0.70 0.66 0.68 0.68 5 2.40 2:34 2.28 2.28 2.30 2.24 2.26 2.24 2.22 2.26 2.62 2.68 2.64 2.66 2.72 2.78 2.76 2.70 2.64 2.58 6 0.78 0.84 0.82 0.76 0.84 0.80 0.76 0.90 0.98 0.84 1.16 1.20 1.16 1.14 1.24 1.26 1.24 1.10 1.22 1.10 7 1.70 1.84 1.76 1.62 1.80 1.74 1.66 1.88'1.80 1.72 2.40 2.38 2.40 2.36 2.48 2.40 2.48 2.44 2.52 2.32 8 1.28 1.36 1.40 1.20 1.40 1.32 1.24 1.37 1.36 1.28 1.52 1.56 1.52 1.50 1.60 1.56 1.56 .148 1.60 1.56 9 1.02 1.08 1.04 1.02 1.10 1.08 1.04 1.10 1.10 1.00 1.08 1.24 1.16 1.10 1.20 1.20 1.20 1.18 1.20 1.24 10 4.78 5.12 5.02 4.40 5.14 4.94 4.70 5.26 5.24 4.84 6.61 6.38 6.24 6.10 6 .5 2 5.42 6.48 6.20 6.54 6.22 11 1.00 1.02 1.06 0.98 0.96 1.04 1.02 1.00 1.02 1.06 1.00 1.02 0.98 0.98 1.04 1.00 0.90 0.98 1.02 1.00 in 12 0.60 0.58 0.60 0.52 0.50 0.54 0.50 0.54 0.52 0.60 0.52 0.50 0.48 0.52 0.52 0.46 0:48 0.52 0.56 0.54 13 0.88 0.94 0.94 0.90 0.86 0.88 0.82 0.90 0.84 0.94 1.06 1.06 1.04 1.02 1.08 1.06 1.06 1.04 1.10 1.04 ' 14 6.58 6.66 6.33 6.23 6.23 6.41 6.16 6.66 6.25 6.58 6.83 7.41 7.33 6.58 7.99 7.66 8.08 6.66 7.58 7.41 15 2.20 2.34 2.20 2.12 2.28 2.28 2.12 2.30 2.32 2.32 2.80 2.84 2.80 2.68 3.00 2.88 2.88 2.72 2.96 2.76 16 1.80 1.90 1.70 1.68 1.90 1.80 1.78 1.88 1.80 1.88 2.32 2.40 2.36 2.24 2.48 2.44 2.36 2.24 2.44 2.24 17 0.40 0.40 0.40 0.38 0.38 0.40 0.38 0.40 0.36 0.40 0.44 0.42 0.44 0.42 0.46 0.48 0.44 0.40 0.46 0.40 18 0.40 0.42 0.42 0.40 0.42 0.42 0.42 0.44 0.44 0.42 0.48 0.44 0.48 0.46 0.48 0.47 0.46 0.38 0.50'0.46 19 0.54 0.54 0.54 0.50 0.58 0.54 0.54 0.58 0.58 0.56 0.56 0.54 0.56 0.56 0.56 0.58 0.56 0.48 0.60 0.54 20 .120 .132 .120 .126 .120 .120 .120 .126 .115 .132 .110 .110 .120 .110 .125 .125 .115 .110 .120 .115 21 1.00 1.10 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.02 0.98 0.98 0.96 1.00 1.08 1.02 0.98 0.96 1.06 0.98 22 1.40 1.42 1.40 1.30 1.32 1.38'1.30 1.30 1.38 1.42 1.26 1.26 1.24 1.20 1.22 1.22 1.24 1.26 1.34 1.24 23 0.76 0.84 0.78 0.76 0.76 0.80 0.72 0.80 0.74 0.80 0.86 0.84 0.86 0.80 0.84 0.90 0.86 0.92 0.88 0.84 24 0.96 1.00 0.98 0.70 0.90 0.70 0.86 0.90 0.86 0.70 1.10 1.14 1.20 1.12 1.20 1.18 1.16 1.20 1.18 1.14 Appendix C - (continued)

Notostira elongata Megaloceraea recticornis 1 8.80 L80 8.74 L72 6.80 6.80 6.78 0.79 0.70 0.74 0.64 0.70 0.64 0.60 0.62 6.64 6.74 6.74 6.72 6.72 2 0.66 0.74 0.76 0.70 0.70 0.74 0.73 0.65 0.68 0.70 0.70 0.70 0.66 0.60 0.70 0.74 0.70 0.72 0.76 0.72 3 0.24 0.26 0.26 0.26 0.24 0.26 0.26 0.24 0.22 0.22 0.60 0.68 0.62 0.66 0.62 0.62 0.62 0.58 0.54 0.62 4 0.42 0.44 0.42 0.40 0.40 0.44 0.42 0.40 0.38 0.40 0.62 0.64 0.64 0.64 0.64 0.68 0.68 0.64 0.68 0.70 5 2.12 2.24 2.18 2.08 2.14 2.24 2.19 2.08 1.98 2.06 2.56 2.72 2.56 2.50 2.58 2.68 2.74 2.68 2.70 2.76 6 1.32 1.32 1.30 1.30 1.30 1.32 1.46 1.32 1.32 1.34 1.28 1.20 1.16 1.36 1.24 1.30 1.22 1.30 1.28 1.26 7 2.44 2.34 2.32 2.30 2.26 2.26 2.92 2.56 2.48 2.60 2.72 2.52 2.44 2.80 2.56 2.72 2.64 2.72 2.66 2.56 8 2.04 1.39 1.84 1.80 1.78 1.78 2.12 2.00 2.00 2.10 3.12 2.88 2.99 2.96 3.00 2.96 2.92 3.04 2.92 2.92 9 1.30 1.28 1.26 1.26 1.30 1.30 1.36 1.36 1.32 1.46 1.16 1.08 1.08 1.10 1.16 1.14 1.20 1.08 10 7.10 6.74 6.82 6.66 6.64 6.66 7.86 7.24 7.12 7.50 8.28 7.68 7.67 7.90 8.04 7.92 8.06 7.82 11 1.04 1.06 1.04 1.02 1.06 1.06 1.10 0.96 0.94 1.06 1.04 0.96 0.90 1.04 0.98 1.00 0.98 1.02 1.00 1.04 in 12 0.64 0.65 0.66 0.64 0.64 0.66 0.71 0.60 0.58 0.60 0.50 0.52 0.50 0.56 0.58 0.62 0.64 0.60 0.60 0.58 13 1.20 1.10 1.14 1.08 1.16 1.18 1.20 1.08 1.06 1.10 1.04 1.02 1.02 1.06 1.06 1.08 1.04 1.12 1.00 1.10 14 7.33 7.66 7.66 7.34 7.24 7.58 7.74 6.66 6.99 7.24 7.41 7.25 7.00 7.75 7.08 8.66 8.50 8.08 7.66 7.33 15 3.04 2.84 2.84 2.92 2.88 2.92 3.36 2.92 2.88 2.92 3.32 3.20 3.08 3.36 3.32 3.40 3.36 3.56 3.36 3.44 16 2.68 2.52 2.52 2.64 2.56 2.60 2.96 2.62 2.62 2.68 2.92 2.72 2.72 3.00 2.84 2.96 2.86 2.96 2.76 2.88 17 0.42 0.40 0.42 0.41 0.41 0.44 0.46 0.33 0.34 0.40 0.36 0.34 0.36 0.36 0.36 0.38 0.40 0.38 0.38 0.42 18 0.54 0.51 0.52 0.53 0.50 0.56 0.60 0.50 0.58 0.56 0.48 0.48 0.38 0.50 0.48 0.48 0.44 0.50 0.50 0.50 19 0.68 0.64 0.66 0.64 0.62 0.66 0.70 0.66 0.66 0.64 0.56 0.54 0.52 0.58 0.52 0.56 0.58 0.58 0.56 0.58 20 .103 .103 .109 .109 .097 .109 .109 .092 .097 .109 .097 .097 .103 .097 .086 .103 .103 .097 .103 .103 21 1.04 1.06 1.02 1.02 1.04 1.08 1.12 0.90 0.92 1.04 1.00 0.96 1.02 1.00 1.00 1.04 1.04 1.04 1.00 1.04 22 1.28 1.30 1.26 1.24 1.28 1.36 1.32 1.20 1.24 1.26 1.26 1.20 1.22 1.26 1.22 1.28 1.46 1.32 1.26 1.36 23 0.80 0.82 0.82 0.92 0.92 0.96 0.89 0.79 0.80 0.79 0.76 0.70 0.82 0.78 0.72 0.84 0.82 0.82 0.76 0.78 24 1.20 1.10 1.00 1.02 1.08 1.14 1.06 1.06 1.06 0.60 1.16 1.10 1.14 1.16 1.14 1.16 1.18 1.18 1.18 1.20 Appendix C - (continued)

Trigonotylus ruficornis Leptopterna dolabrata a 1 0.38 0.40 0.38 0.36 0.38 L38 0.38 0.38 0.40 0.34 0.68 0.72 0.60 6.72 0.66 0.68 0.56 0.62 2 0.30 0.30 0.32 0.38 0.40 0.40 0.40 0.36 0.42 0.37 0.88 0.80 0.80 0.76 0.72 0.68 0.62 0.68 0.68 3 0.30 0.30 0.36 0.36 0.40 0.40 0.38 0.36 0.40 0.30 0.56 0.52 0.56 0.56 0.48 0.54 0.56 0.56 0.56 0.48 4 0.30 0.36 0.38 0.38 0.38 0.38 0.40 0.38 0.40 0.38 0.64 0.66 0.64 0.62 0.60 0.60 0.60 0.62 0.62 0.60 5 1.28 1.36 1.44 1.48 1.56 1.56 1.56 1.48 1.22 1.29 2.76 2.70 2.60 2.66 2.46 2.46 2.42 2.38 6 0.52 0.56 0.60 0.56 0.56 0.56 0.64 0.62 0.60 0.58 1.06 1.02 1.02 1.04 0.96 0.98 1.02 1.02 1.02 1.00 7 1.28 1.56 1.44 1.44 1.48 1.42 1.60 1.56 1.60 1.40 2.60 2.64 2.58 2.56 2.44 2.56 2.72 2.68 2.60 2.40 8 1.40 1.58 1.48 1.52 1.46 1.42 1.60 1.54 1.56 1.40 1.68 1.80 1.70 1.76 1.64 1.72-1.76 1.68 1.72 1.68 9 0.52 0.60 0.60 0.54 0.52 0.56 0.62 0,58 0.58 0.58 1.04 1.00 1.04 1.00 1.02 1.02 1.04 1.08 1.04 10 3.72 4.30 4.12 4.06 4.02 3.96 4.46 4.30 4.34 3.96 6.38 6.46 6.26 6.36 6.28 6.52 6.42 6.42 6.12 11 0.70 0.74 0.80 0.76 0.78 0.76 0.78 0.80 0.82 0.80 1,24 1.22 1.24 1.18 1.12 1.18 1.18 1.20 1.16 1.12 ul 12 0.42 0.40 0.44 0.42 0.42 0.40 0.40 0.40 0.42 0.44 0.68 0.68 0.72 0.68 0.58 0.60 0.62 0.64 0.56 0.64 13 0.72 0.74 0.78 0.76 0.82 0.74 0.80 0.80 0.84 0.82 1.10 1.10 1.08 1.06 0.98 1.16 1.12 1.18 1.02 0.98 ' 14 4.16 4.16 4.40 4.12 4.20 4.44 4.44 4.72 4.68 4.60 7.58 6.46 6.33 6.91 6.16 6.16 6.75 7.50 7.33 7.08 15 1.60 1.76 1.04 1.70 1.72 1.78 1.88 1.82 1.90 1.84 2.88 2.88 2.80 2.84 2.72 2.90 2.84 2.80 2.76 2.68 16 1.30 1.36 1.48 1.42 1.46 1.42 1.56 1.50 1.50 1.42 2.30 2.28 2.24 2.20 2.08 2.32 2.32 2.20 2.12 2.08 17 0.24 0.26 0.28 0.26 0.30 0.32 0.30.0.30 0.30 0.30 0.50 0.44 0.48 0.48 0.38 0.44 0.46 0.44 0.40 0.36 18 0.28 0.26 0.30 0.28 0.26 0.30 0.32 0.30 0.30 0.32 0.48 0.49 0.48 0.48 0.46 0.50 0.48 0.46 0.46 0.44 19 0.38 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.40 0.42 0.64 0.64 0.60 0.64 0.60 0.66 0.66 0.58 0.62 0.60 20 .06; .090 .080 .074 .086 .080 .086 .090 .090 .074 0.15 0.12 0.13 0.13 0.11 0.14 0.13 0.12 0.12 0.12 21 0.70 0.74 0.76 0.78 0.80 0.82 0.82 0.82 0.88 0.82 1.28 1.12 1.20 1.14 1.12 1.14 1.18 1.22 1.10 1.10 22 0.83 0.94 0.98 0.96 1.02 1.04 0.98 1.00 1.02 0.96 1.28 1.46 1.48 1.48 1.46 1.50 1.58 1.58 1.44 1.50 '23 0.46 0.50 0.50 0.46 0.48 0.54 0.56 0.50 0.56 0.44 0.84 0.80 0.84 0.80 0.80 0.82 0.84 0.80 0.84 0.82 24 0.78 0.80 0.96 0.84 0.96 0.80 0.86 0.86 0.76 0.82 0.50 0.54 0.42 0.32 1.04 1.22 1.12 0.64 0.60 1.12

Appendix C - (continued)

Leptopterna ferrugata Teratocoris antennatus

1 0.48 0.60 0.60 0.60 0.66 0.56 0.54 0.64 0.52 0.56 0.24 0.34 0.36 a 0.32 0.32 2 0.56 0.62 0.63 0.62 0.72 0.56 0.60 0.60 0.50 0.52 0.30 0.26 0.30 0.24 0.31 3 0.60 0.56 0.56 0.50 0.63 0.52 0.54 0.70 0.56 0.60 0.26 0.23 0.28 0.26 0.24 4 0.60 0.60 0.60 0.70 0.64 0.58 0.58 0.60 0.52 0.60 0.36 0.36 0.34 0.32 0.30 5 2.24 2.38 2.39 2.42 2.65 2.22 2.26 2.54 2.04 2.28 1.16 1.19 1.28 1.13 1.14 1.17 6 0.90 1.00 0.92 0.96 0.94 0.84 0.92 1.00 0.88 0.90 0.80 0.80 0.82 0.74 0.78 0.72 7 2.10 2.16 1.83 2.20 2.14 1.92 2.00 2.10 1.96 2.10 1.38 1.30 1.32 1.22 1.18 1.26 8 1.44 1.42 1.40 1.48 1.50 1.48 1.46 1.52 1.54 1.66 1.20 1.18 1.20 9 0.80 0.82 0.80 0.90 0.84 0.80 0.90 0.80 0.80 1.00 I 10 5.24 5 .3 8 4.95 5.54 5.42 5.04 5.28 5.18 5.46 4.38 N. 11 1.14 1.04 1.08 1.08 1.10 1.04 1.10 1.06 0.90 1.08 0.84 0.84 0.84 0.74 0.78 0.74 12 0.60 0.56 0.60 0.64 0.64 0.48 0.52 0.56 0.46 0.54 0.42 0.41 0.44 0.36 0.36 0.37 13 1.16 0.98 1.16 0.98 1.14 0.96 0.98 1.02 0.92 1.00 0.66 0.70 0.6.9 0.58 0.60 0.60 14 7.08 6.66 6.91 6.74 6.74 6.91 6.66 7.16 5.83 7.16 4.72 4.00 5.08 4.08 4.12 3.72 15 2.32 2.36 2.10 2.30 2.56 2.08 2.20 2.56 2.08 2.40 1.60 1.62 1.60 1.40 1.56 1.34 16 1.88 1.88 1.70 1.90 2.08 1.88 1.90 2.02 1.70 2.10 1.48 1.50 1.46 1.30 1.49 1.24 17 0.48 0.48 0.46 0.48 0.46 0.38 0.44 0.42 0.34 0.38 0.27 0.26 0.26 0.25 0.26 0.24 18 0.42 0.46 0.38 0.44 0.48 0.44 0.46 0.50 0.42 0.46 0.26 0.26 0.28 0.24 0.24 0.24 19 0.54 0.54 0.50 0.54 0.58 0.52 0.60 0.60 0.50 0.50 0.42 0.42 0.42 0.40 0.40 0.36 20 .097 .109 .097 .109 .097 .092 .097 .098 .080 .086 .080 .086 .086 .074 .074 .074 21 1.20 1.10 1.18 1.12 1.10 1.02 1:.04 1.04 0.94 1.04 0.76 0.76 0.78 0.68 0.68 0.68 22 1.44 1.42 1.44 1.44 1.44 1.42 1.40 1.52 1.16 1.38 0.98 1.01 0.98 0.80 0.88 0.80 23 0.84 0.74 0.72 0.82 0.84 0.76 0.76 0.90 0.64 0.72 0.58 0.62 0.60 0.56 0.52 0.51 24 0.34 0.46 0.40 0.36 0.34 1.08 1.06 1.10 0.60 0.74 0.52 0.68 0.50 0.40 0.40 0.42

Appendix C - (continued)

Teratocoris caricis Teratocoris viridis Teratocoris saundersi 1 0.30 .320 0.30 0.30 0.30 0.33 320. 0.32 0.30 0.35 0.34 0.30 0.30 0.31 0.30 0.30 2 0.32 0.30 0.30 0.30 0.31 0.32 0.28 0.30 0.30 0.30 0.32 0.29 0.30 0.29 0.30 0.32 3 0.30 0.30 0.30 0.30 0.32 0.30 0.30 0.30 0.32 0.32 0.30 0.30 0.30 0.30 0.30 0.29 4 0.36 0.39 0.34 0.38 0.36 0.36 0.34 0.35 0.36 0.38 0.34 0.34 0.34 0.36 0.35 0.36 5 1.26 1.31 1.24 1.28 1.28 1.31 1.24 1.27 1.28 1.35 1.30 1.23 1.24 1.26 1.25 1.25 6 0.96 0.92 0.88 0.86 0.90 0.92 0.84 0.88 0.88 0.78 0.74 0.78 0.88 0.90 0.88 0.96 7 1.90 1.84 1.76 1.66 1.80 1.90 1.68 1.80 1.54 1.42 1.42 1.42 1.86 1.82 1.76 1.86 8 1.24 1.30 1.24 1.20 1.20 1.26 1.16 1.22 1.10 1.08 1.06 1.03 1.32 1.36 1.22 1.26 9 1.08 1.04 1.08 1.06 0.92 0.96 0.94 0.89 1.08 1.10 1.06 1.11 10 5.18 5.12 4.74 4.96 4.44 4.24 4.16 4.12 5.14 5.18 4.92 5.19 11 0.84 0.94 0.84 0.92 0.86 0.86 0.87 0.84 0.91 0.92 0.88 0.83 0.82 0.90 0.84 0.84 in 12 0.48 0.56 0.48 0.48 0.46 0.48 0.56 0.49 0.46 0.48 0.48 0.48 0.46 0.50 0.48 0.43 ' 13 0.66 0.68 0.62 0.70 0.66 0.64 0.68 0.66 0.73 0.72 0.68 0.70 0.62 0.68 0.64 0.68 14 4.52 4.44 4.60 4.60 4.80 4.60 4.80 4.78 4.74 4.34 4.40 4.52 4.40 4.76 4.56 4.72 15 2.00 2.02 1.88 1.90 1.90 1.90 1.74 1.90 1.82 1.80 1.70 1.66 1.90 2.00 1.90 1.94 16 1.88 1.94 1.82 1.76 1.80 1.88 1.66 1.82 1.83 1.70 1.62 1.60 1.76 1.90 1.80 1.80 17 0.30 0.36 0.29 0.37 0.32 0.32 0.32 0.30 0.32 0.34 0.33 0.32 0.32 0.32 0.29 0.29 18 0.36 0.37 0.36 0.34 0.32 0.36 0.34 0.38 0.36 0.32 0.33 0.32 0.36 0.36 0.34 0.35 19 0.50 0.50 0.48 0.50 0.49 0.52 0.47 0.50 0.48 0.44 0.45 0.46 0.52 0.50 0.50 0.48 20 .086 .090 .103 .097 .103 .092 .103 .095 .097 .097 .097 .092 .097 .097 .097 .097 21 0.76 0.80 0.80 0.87 0.76 0.76 0.80 0.76 0.84 0.84 0.80 0.78 0.74 0.80 0.78 0.80 22 1.04 1.07 1.02 1.07 1.02 1.02 0.98 0.98 1.09 1.06 1.02 0.96 0.96 1.00 0.98 0..96 23 0.60 0.64 0.60 0.60 0.60 0.60 0.58 0.60 0.66 0.62 0.62 0.58 0.59 0.60 0.56 0.58 24 0.82 0.68 0.80 0.54 0.76 0.76 0.60 0.78 0.82 0.60 0.60 0.60 0.56 0.52 0.60 0.60 - 459 - Appendix D - Changes in county designations brought about by the Local Government Act of 1972.

Place name Old county New county or region Aviemore Inverness Highland Region Bodesbeck Dumfs Dumf.& Galloway Region Bonar Bridge Sutherland Highland Region Bournemouth Hants Dorset Brigsteer Westm Cumbria Devil's Gallop Lanes Cumbria Edinburgh Midlothian Lothian Region Hampton Court Mddsx Greater London Hounslow Mddsx Greater London Kinloch Rannoch Perths Tayside Region Leadhills Dumfs Strathclyde Region Longashton Som Avon Longsleddale Westm Cumbria Malham Tarn Yorks N. Yorks Merlewood Lanes Cumbria Roudsea Lanes Cumbria Rusland Moss Lanes Cumbria Rye Sussex E. Sussex Scratchwood Mddsx Greater London Slough Bucks Berks Ubley Som Avon Wanlockhead Dumfs Dumf. & Galloway Region - 460 -

Acknowledgements

I should like to record here my debt of gratitude to the following people who have given me much help and encourage- ment during the last three years: to Professor T.R.E. Southwood, my Imperial College director of studies, for provision of nymphal material and much advice; to Dr W.J. Knight, my British Museum supervisor, for discussions and continued support; to Dr P. Freeman and the Trustees of the British Museum (Natural History) for financial support and use of extensive facilities; to Bill Dolling for sharing his knowledge of the Heteroptera; to members of the EM Unit and Photographic Studio, especially Don Claugher for advice on electron microscopy, and Frank Greenaway for Cibachrome prints; to Mr R.G. Davis for computing and advice on numeri- cal taxonomy; to Dave Cooper for histology; to the persons mentioned in the list of collectors (Appendix A); to the Entomology Library staff, especially Pam Gilbert; and final- ly, but not least, to Dr Maggie Manson for typing, proof- reading and moral support. - 461 -

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COOMBS, C.W. & HALSTEAD, D.G.H (1976) J. stored Prod. Res., 12, 121-128. This reference gives a complete bibliography of the published works of G.E. Woodroffe who wrote widely on - Heteroptera.