(HOMOPTERA: COCCOIDEA) by JICHANI

MORPHOLOGY AND TAXONOMY CF ADULT MALES .OF THE FAMILY CCCCIDAE (HOMOPTERA: COCCOIDEA) by JICHANI\'ES HUMAN GILIGMEE M.Sc. (Agric.) Thesis submitted for the degree of Doctor of Philosophy in the University of London. Dept. of Zoology EL Applied Entomology, Imperial College of Science & Technology, South Kensington, London, S.W.7. June, 1964. ABSTRACT The males of 23 species (representing 19 genera) of the family Coccidae have been described and illustrated in detail and a general account of the external morphology of male Coccidae is given. A number of structures present in other male Coccoidea but not hitherto observed in the Coccidae have been recorded. The relationships of the lecanoid type of male with the margaroid and diaspidoid types have been discussed and the males of two families of the lecanoid type (Coccidae and Pseudococcidae) have been compared with each other. Within the Coccidae the males were often found to reveal different relationships from the female and a classification is suggested which differs from the classifications based on female characters. The results of this study is in accor- dance with recent discoveries that the characters of the male are valid at all taxonomic levels, including genera and species. Detailed keys to groups of genera, genera and species have been provided. 2 CONTENTS Page: ACKNOWLEDGMENTS • • • • . 6 I. INTRODUCTION AND REVIEW OF THE LITERATURE 8 (a) Introduction . • • . 8 (b) Review of the Literature • II II. MATERIAL AND TECHNIQUE • • • 15 III. GENERAL MORPHOLOGY 19 A. General Characteristics • 20 B. The Head 22 1. Head Capsule • • • 22 2. Antennae • • . 30 C. The Thorax 33 1. Prothorax . • • . 33 2. Mesothorax • • 37 3. Metathorax • • • 46 4. Wings and Halteres . 50 5. Legs . • • 52 D. The Abdomen 55 (a) The pregenital segments 56 (b) The genital segment and external genitalia • • 61 IV. DESCRIPTION OF THE SPECIES 63 Eulecanium Group 63 1. Eulecanium tiliae (L) . 63 2. Nemolecanium abietis Borchs. 70 3. Phys.kermes piceae (Schr.). 77 4. Rhodococcus spiraeae (Borchs.) 83 5. Palaeolecanium bituberculatum (T.T.) . 89 6. Phyllostroma myrtilli (Kalt.) 96 7. Filippia viburni (Sign.) 105 8. •Ctenochiton sp . 114 9. Ericerus pela (Chay.) 123 10. Genus A . • • . 11. Sphaerolecanium prunastri (Fonsc.) • • • Eriopeltis group 150 12. Eriopeltis festucae (Fonsc.) 150 13. Eriopeltis sp. 160 14. Luzulaspis luzulae (Dufour) 170 .4 Inglisia group 180 15. Inglisia theobromae Newst. 180 Coccus group 190 16. Coccus hesperidum L. 191 17. Genus B (near Pulvinaria) spa 201 Pulvinaria betulae (L.) 209 19. Pulvinaria acericola (Walsh & Riley). 218 20. Parthenolecanium corni (Bouche) 226 21. Parthen•lecaium pomeranicum (Kaw.) • 237 22. Ceroplastes berliniae Hall . 245 23. Ceroplastes sp.. • • 253 V. DISCUSSION 261 (a) Taxonomic Significance of the Characters • . 264 (b) Classification and Interrelationships of the Coccidae based on the Males 284 (i) Groups of Genera . 284 (ii) Geritera . • 301 ( -LI) S • • • 311 (c) Relationships of the Coccidae with •her subdivisions of the Coccoidea 311 VI. KEYS . 224 VII. SUMMARY . 343 VIII. REFERENCES . 3,4.7 IX. LIST OF FIGURES . 353., X. EXPLANATION OF FIGURES . 354 XI. LIST OF ABBREVIATIONS . 355 XII. FIGURES . 359 6 ACKNOWLEDGMENTS I would like to express my gratitude to Dr. K. Boratynski who supervised the work, Mr. R.G. Davies for advice and constructive criticism and Prof. O.V. Richards, F.R.S., for providing facilities at the Imperial College of Science and Technology. I wish to thank Dr. D.J. Williams of the Commonwealth Institute, London, and Mr. G. de Lotto of the Research Institute for Plant Protection, Pretoria who supplied and identified some of the material, and also the large number of other workers who so generously made material avail- able for this study. They are: Prof. N.S. Borchsenius, Zoological Institute, Academy of Sciences, LeninL;rad; Dr. J.M. Cherret, University College of North Wales, Bangor, Wales; Dr. M.S.K. Ghauri, Commonwealth Institute of Entomology, London; Prof. M. Kosztarab, Virginia Polytechnic Institute, Blacksburg, Virginia; Prof. Z. Kawecki, Agricultural University, Warsaw; Dr. G. Matesova, Insti- tute of Zoology, Academy of Sciences, Alma-Ata, Kazakhstan; Ing. MitiC-Muiina, Institute for Plant Protection, BelLrade; Mr. J. Munting, Plant Protection Research Institute, Pretoria; Dr. J. Rehagek, Viroloical Institute, Bratislava; Dr. J.G. Theron, University of Stellenbosch, Stellenbosch. I am also indebted to the Trustees of the British Museum (Nat. Hist.) for permission to study material in their collection. Finally, I wish to thank the authorities of the Commonwealth Scholarship and Fellowship Plan for financial assistance, and the University of Stellenbosch and the Government of the Republic of 7 South Africa for granting me leave to undertake this study. 8 I INTRODUCTION AND REVIEW OF THE LITERATURE (a) INTRODUCTION The economic importance of the Coccoidea attracted the attention of many research workers to this superfamily and numerous detailed studies of various aspects of its morphology, internal anatomy, taxonomy, bionomics and also Genetics have been carried out. With the exception of the Genetical studies most of the work has been concerned only with the adult females while the males have received very little attention. Conforming to the treatment of other insects, some of the earlier workers like Targioni-Tozetti (1867), Putnam (1879) and Berlese (1893, 1896) tried to give equal attention to both sexes, but soon afterwards interest in the males declined and reference to them consisted only of brief, superficial descriptions which contributed little to the already existing knowledge. As already pointed out and discussed by Theron (1958), Beardsley (1960) and Ghavri (1962) this situation arose mainly because of the biological peculiarities of the superfamily, which render the males very difficult to obtain, and because the general characteristics of the male make morphological studies technically difficult. It was also commonly accepted, perhaps for the s6.he of convenience, that the males - which apart from the absence of functioning mouth parts are "normal" insects - were of but little taxonomic value. Consequently the identifica- tion and classification of the Coccoidea were, and still are, based mainly on the characters of the comparatively large and easily obtainable females. These females afford many characters which can be used to separate species and groups of species but it is very unsatisfactory to determine phylogeny and interrelationships on the basis of their characters alone - not so much because they are neotenic, highly specialised and more or 9 less degenerate, but because one is using only a fraction of the potential information. Not surprisingly therefore, considerable disagreement and confusion have arisen regarding the affinities of larger or smaller groups within the superfamily. Morrison (1928) in a paper on the classification of the Margarodidae, was one of the first to question the reliability of a system of classification based on the characters of the adult female alone. He also studied stages other than the adult female and found that the limits of any association of genera based on the characters of one stage disagreed from that based on the characters of any other stage. He made the interesting discovery that "the adult male stage of any species possesses morphological characters that appear to be of marked value in indicating the broader relationships of the genus to which it may belong", and he compiled keys to separate the adult males even within genera (e.g. Drosicha). The fact that the male was of importance in distinguishing lower taxonomic categories was also illustrated by the work of Stickney (1934a), who constructed keys to the adult males of the species that he included in the Phoenicococcinae. Subsequently the necessity to study the males gradually began to be realised and Ferris (1942, 1950, 1957) repeatedly stressed the need for detailed studies of the male. Ferris (1937, 1957) and also Balachowsky (1937) were of the opinion that a knowledge of the morphology of the male would only be of importance in determining relationships of the higher c?_terry,:sies, but of little value on the specific level. As a result of the revival of interest in the males a series of papers dealing with this sex alone was published (Jancke, 1955; Ezzat, 1956; MIstshik, 1958); the male also received more attention in general works (Borchsenius, 1957) and in the descriptions of new species (Hadzibejli, 1955). I0 Although these studies were lacking in detail and proper inter- pretation of the morphology, they seemed to support the view that males could be used to separate lower categories as well. A major advance was made when Theron (1958)carried out a detailed study of the general morphology of the males of 7 genera, representing 4 families. He suggested homologies for the various structures and indicated the relationship of these families with each other and the other Sternorrhyncha, thus laying a firm foundation on which future studies could be built. This work was followed up by detailed studies of the males of 3 species of the Pseudococcidae (Gdliomee, 1961) and 24 species of the Diaspididae (Ghauri, 1962), while Beardsley (1960, 1962) published brief descriptions and notes on 35 species of the Pseudococcidae. In the Diaspididae, which is probably the best known of all the Coccoidea, a number of subdivisions (subfamilies, tribes, subtribes and groups) have been recognised on the

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