BOOK NUMBER % k § z § - p § M S Theses R t o n The Balme L lh ra ru 3 0692 1078 6861 2 University of Ghana http://ugspace.ug.edu.gh KARYUTYPi^B OF 5 0HB GilitNAlAH SHIejuD-BUCio IHB HIGHER SYbTeHAl'ICa OF (Ha& Ut'lli^A : By KWAjyJB aFRMI ifUAMH, B .B c . (HUnuURB) A THjSBIB bUBiiITicii) FOR iH u i> u ^ u ^ OF Mn^ThR OF a a t m e u iavjiixioiTY o f g^aW a, L eajun , 19 8 0 . SriPTiui-lBeR, 1980 University of Ghana http://ugspace.ug.edu.gh DJSPAKT^WT OF ZiOOLUUY UNIVi^ITY OF uKitNA LtitiGN GHiiNA DEGLAiiAllOh This is to certify that this thesis has not been submitted for a degree to any other University. It is entirely my own work, and all help has been duly acknowledged. University of Ghana http://ugspace.ug.edu.gh DiiDICiiTION This work is dedicated to Him, through whom, and of vftiom are all things, the Lord God Almighty, uho makes all things possible; to Grace and my Parents, for their ever present love; and to Professor R. Kumar for his valuable contributions to Lntomology in Ghana. University of Ghana http://ugspace.ug.edu.gh My sincere gratitude goes to Professor k. Kumar, ray Supervisor and head of the department of zoology, University of Ghana Legon, for his keen interest in this work, for his critical guidance and suggestions and for reading through the work* I wish to thank also Mr P.U.O* Kpordugbe for valuable assistance with the technical aspects of the project, and to Professor w.^ . Coker for reading through the manuscript. I am indebted to Mr M. Adjo (driver), to i1. Ansah, Mr B. j-.remong and the other departmental assistants for assistance in diverse ways. My thanks are also due to the uirector, Commonwealth Institute of Entomology for confirming the identity of insects determined by Prof. K* Kumar. The work was supported by a bursary from uhana government and a grant from the Cocoa research Co-ordinating Committee, Legon. finally, I am highly indebted to Mr. N# Owusu, Zoology Department, University of G-hana, Legon for his diligent and meticulous typing of this study. University of Ghana http://ugspace.ug.edu.gh Data on 2n-numbers of some fifty-five (55) species , belonging to six families of some Ghanaian shield bugs, collected in Southern Ghana, west Africa are presented. Microphotographs and tracings of karyotypes observed for each species are produced as figures. Histograms of chromosome numbers of the various families have been constructed and these results together with the existing information in literature, is analyzed and discussed. In all two hundred and seventeen (217) species of Pentatomoidea were found to be known cytologically in the literature. .*ith the present work the total number of species of shield-bugs where the Karyotypes is now known stands at two hundred ana sixty-six (266). The following karyotype groups appear to emerge from the study; 14 and 12. Under the group '\k is tne family Pentatomidae, whilst the families, Scutelleridae, Gydnidae and Plataspiaae fall under the karyotype group 12. i\lo basic chromosome number can however be cited for the other families of Pentatomoidea, namely, i/inidoridae, Tessaratomidae, Acanthosomatidae and Urostyliaae, because the cytological data for each of these are too scanty, for any such conclusions to be arrived at. The inter-relation of various groups in the superfamily Pentatomoidea are discussed, and the present investigations tend to support tne work ana conclusions arrived at by morpho-taxonomical studies. abstract University of Ghana http://ugspace.ug.edu.gh TaBLL OF CQI)TiklVl'b PaG& Title 1 Declaration 11 Dedication ii;L Acknowledgement 1V Abstract v 'Table of contents V1 1 General Introduction 1 2 Materials and Methods 7 3 Results 3.1 Introduction 3.2 Meiosis 10 3.3 Description of meiosis 13 3*3*1 Acrosternum heegeri Fieb. 1^ 3*3*2 Aeliomorpha sp.? graseoflava otRl 13 3.3*3 Aeptus singularis (Dallas) 16 3*3*^ Aethemenes chloris (rtestw.) 17 3«3*5 Aspavia acuminata (Mont.) l8 3.3.6 Aspavia armigera (Fabr.) 19 3.3.7 Aspavia hastator (Fabr.) 20 SECTION University of Ghana http://ugspace.ug.edu.gh Page 21 22 23 24 25 2? 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 Atelocera serrata (Fabr• ) Bathycoelia thalassina (H*-o) Bathycoelia rodhaini (Gchout.) Benia sp. A Carbula capito StSl Carbula carbula (Dist.) Carbula marginella bt§l Carbula melacantha stal Garbula sp* nr. s.jostedti (ochout.) Gaura pugillator (Fabr*) Diploxys bipunctata (Amyot et Serville) Amaxosana punctata Distant Durmia haedula btal Purmia lutulenta stSl Durmia sp* Dymantis grisea Jen-Haar Lysarcoris inconspicuus (n.-o) Halyomorpha reflexa (sign) Leri da punctata J^r’alisot iieauvois) Macrorhaphis acuta Dallas Nezara viridula (Linn.) University of Ghana http://ugspace.ug.edu.gh SECTION PAGE 3.3.29 Nezara viridula var smaragdula (iabr.) **3 3.3.30 Nezara viridula var torquata (Jruton) 44 3.3.31 Antestia sp. 5^ 3.3.32 Antestia sp. s.l 3.3.33 Antestia sp.? immunda 47 3.3.34 Antestiopsis sp.s.l 48 3.3.35 Piezodorus hybneri (unelin) 49 3.3.36 Scotinophara fibulata (bchout.) 50 3.3.37 bepontia misella ot§l 51 3.3.38 i>oma verrucosa (hont.) 52 3.3.39 Veterna sanguineirostris btal 53 3.3.40 Macrina .iuvenca (Burm.) 55 3-3.41 Callidea duodecimpunctata stfil 56 3.3.42 Hotea subfasciata (Westw.) 57 3.3.43 Sphaerocoris testudogrisea st§l 58 3.3.44 Ste&anocerus multipunctata (de Geer) 59 3.3.45 Brachyplatys incertus (walker) 60 3.3.46 Brachyplatys testudonigra stal 61 3.3.47 Coptosoma nubila Germ. 62 3.3.48 Coptosoma stali (Mont.) 63 University of Ghana http://ugspace.ug.edu.gh Piitiiii 6k 65 66 67 67 68 68 71 73 74 77 78 79 83 86 97 Coridius cuprifer (Westw.) Piezosternum calidum btal Macroscytus sp. Discussion bystematics Karyotypes and classification Family Pentatomidae Family bcutelleridae Family Brachyplatidae (Plataspidae) Dinidoridae tessaratomidae Family Gydnidae Other families of Pentatomoidea bummary Literature cited Appendix tables University of Ghana http://ugspace.ug.edu.gh SECTION 1 GENERAL INTRODUCTION A significant advance in our knowledge of the higher classification of the terrestrial Heteroptera (Geocorisae) was made by leston, Pendergrast and Southwood (1954) who, following evans* (19^) precedent in Homoptera, divided Geocorisae into two groups, namely Pentatomorpha and Cimicomorpha. The grouping was based on evidence from external male genitalia, trichobothria, wings, spermatheca, eggs and the salivary glands. Recently, support in favour of this grouping has been furnished by Southwood (1956, eggs), Pendergrast (1957, internal reproductive organs), Scudder (I959i ovipositor), Miyamoto (1961, alimentary organs) and Cobben (1968, 1978 eggs and mouthparts). Thus now it is well acknowledged that the Pentatomorpha and the Cimicomorpha are natural groups. Objections of China (1955)/ regarding the basing of taxonomic categrories on the internal organs have been well countered by iiiyamoto (1961), who rightly contends that the internal structure such as alimentary organs "are one of the basic structures for life and comparatively stable in character11. Within the groups Pentabtomorpha and Cimicomorpha, the relationships of various taxa have been the subject of a good deal of attention and characters like the genitalia and alimentary organs have been studied at some length. Despite these studies, wiue lacunae exist in our knowledge of these characters in various groups of Heteroptera. Pendergrast (1957) in his detailed survey of the reproductive organs University of Ghana http://ugspace.ug.edu.gh of Heteroptera, listed no fewer than eighteen families in males and eight families in females where the knowledge of these organs is totally lacking. In the rest of the families the position at subfamily and tribal level is not satisfactory, e.g in large families such as the Scutelleridae, the internal male reproductive organs are known only in one species. ‘Though a good deal of work has not been done on the higher classification of the Pentatomoidea, the statements by Pendergrast (1957) that, "there is more disagreement over the family and Subfamily groupings of the Pentatomoidea than in the whole complex of families in the Heteroptera*1 still holds (vide infra). The systematics of the group is in a very confused state, and as observed by Kumar (1965)$ "despite our rapidly accumulating information on the families of the Pentatomoidea, we still have a very imperfect understanding of the relationships of various taxa in this vast and complex group11. In recent years, considerable work has been done on the higher classification of Pentatomoidea, particularly on the male genitalia particularly the aedeagi in an expanded condition, structure and function of the ejaculatory reservoir, ovipositor, internal reproductive organs and structure of eggs and mouthparts. among such workers are, Leston (195^ b, 1955» 1958); Southwood (1956); Pendergrast (1957); Scudder (1959); Kumar (1962, 1965, 1968b); and Gobben (1968, 1978). Inspite of this it is still realised that significant contributions remain to be made in this area, and the need for further work in this field cannot be over-emphasized. As to nov/, taxonomists are increasingly realising that internal University of Ghana http://ugspace.ug.edu.gh organs are less variable than external characters, and may thus at l@stst in some groups serve as valuable phylogenetic indicators (Louis and Kumar 1972). fenna (1951) strongly encourages cytologists interested in a synthesis of cytological and morphological data to gather detailed information about chromosomes and their behaviour, of many more species of animals and plants, and this is exactly what this project addresses itself to do. Chromosome cytology has contributed to insect systematics in several different ways which are not always appreciated very clearly by non-cytologists. ‘Though, it cannot be said to be an automatic solution to all systematic problems, in suitable cases, it can provide critical evidence of a unique kind. Cytologists axe, among other things, concerned with the cytotaxonomic differences which exist between species. These, only rarely involve major differences in the genetic mechanisms, but often consist of differences in chromosome number, and in their sizes and shapes. According to white (1956), such differences may sometimes be used to distinguish 1 •sibling11 or cryptic species that cannot be separated at all, or only with difficulty or uncertainty, on external characters. These are the results of chromosomal rearrangements which have arisen spontaneously ana established themselves in phylogeny. One type of rearrangement may lead to a dimunition in chromosome numbers, while another may produce an increase. The knowledge of how the major differences in genetic mechanisms had arisen in the course of evolution, could be used in establishing University of Ghana http://ugspace.ug.edu.gh the relationships of the insect orders and other 'higher categories1 on a firmer foundation, within a particular species the size range of the chromosomes of a haploid groups is not great, the smallest, usually being no more than one third to half that of the largest. Because, such nuclear cytology provides an independent set of data, which can be critically correlated, and evaluated with the usual morphological findings, it is assuming more ana more importance, though it is still in its infancy. However, as rightly pointed out by Schaefer (1964); cytological work is not easy to evaluate, and the conclusions drawn by some authors discredit the accuracy of their work. Commenting on this Cobben (1968) stressed "that detailed knowledge of cytology and ways of recognizing the different chromosomes, and studies on their behaviour are needed". According to him, unless this is done, attempts to apply "chromosome studies to the solution of problems of major relationships, would be a sheer guess work, such that the results would also be contradictory to current classification, especially, when the cytologist is not a taxonomic specialist of the group in question"* He cited the work of Manna (1958) as an example of how such problems, have arisen, where extremely remote groups are linked together. Actually, such are some of the dangers one runs into, in trying to apply cytology in systematics, but this is to be expected, for such problems cannot be wholly avoided in a work of this nature. It is hoped that with time and more work, such difficulties would be overcome, University of Ghana http://ugspace.ug.edu.gh and the actual advantage of cytology in insect systematics, would be seen and accepted, to provide a unique tool in tackling some of systematics problems as existing at present# Since the time of Montgomery (1901, 1905) Wilson (1911)t the Heteroptera have been undergoing extensive cytological investigations# The main investigations £0 far have been the use of cytological data along with morphological characters to: (a) evaluate the broad mechanical principle underlying changes in karyotypes, (b) evaluate supergenetic, generic and specific classification, liven though among the different groups of Heteroptera, the Pent&tomidae is best known cytologically, (Schachow, 1932; Schrader, 19^ 5&i 19 5^^ , I960, Manna, 1950* 1951* 1958; Leston 195&* etc.)# Generally, though cytology as an additional tool in the field of taxonomy is undeniable, its use is only possible, according to Manna (1958), when we have adequate data at hand, obtained in a planned way of study. Though the Heteroptera are one of the groups of insects that have been the subject of extensive cytological investigation, since the beginning of this century, not all families are known cytologically. Usually, as confirmed by this study, the data are not equally extensive in all families, and in certain cases only a few species have been examined# In the present study, cytological data on fifty-five (55) species belonging to six families of the superfamily Pentatomoidea is presented# All published information on the karyotypes of the Pentatomoidea is University of Ghana http://ugspace.ug.edu.gh reviewed, and interrelationship within the super family are examined* University of Ghana http://ugspace.ug.edu.gh SECTION 2 MATJ^ilAlaD AND MKTHUito Adult male insects of the group Pentatomoidea which constituted the material for the present investigation, were collected locally from within a radius of seventy-five (75) miles of the University campus, in the -bouth-Bastern cocoa growing areas of uhana, west Africa, from February 1979 to m y 1980. The various insects were collected by use of ultra-violet light traps, using mercury bulbs in the night, butterfly and sweeping nets, and also by hand picicing from walls under electric lights, ana on leaves of plants, throughout the period of work. At each particular time of collection, more than one method of collection was used as the need arose, The insects soon after their capture were transported alive to the laboratory and dissected under tap water, and their testis directly transferred to the fixative, namely the Uarnoyfs fixative made up to three (3) parts of isopropanol and one part of glacial acetic acid. These were kept in tubes in an air-conditioned room (average room temperature = 22°C) until squashing. Squashes of testis with Lacto-aceto-orceintain (Warren et al I960) were made. The Lacto-aceto-orcein stain was made up of thirty millilitres (30 mis) each of the following: distilled water, glacial acetic acid, and lactic acid, plus 2 grams of synthetic orcein, heated gently over water bath for about half an hour to one-hour with regular stirring, after which time tne stain was filtered, whilst hot. University of Ghana http://ugspace.ug.edu.gh Chromosome counts and other observations of the squashes were made with the use of a Keichert binocular optical compound microscope with xlO ocular, t^Ox and lOOx(oil) objectives against a green filter. The number of male specimens of each species dissected for testis material varied according to the availability of the particular species. Efforts were made to collect as many and varied species of the male insects, but since their catch depended on their abundance and availability, limitations were imposed on the numbers. In some cases, such as the Bathycoelia rodhaini bchout only one male specimen was caught and dissected, wnilst in others for example, Aspayia armigera (Fabr.) and Sepentia misella btal as many as fifty male specimens were dissected for squashing. Microphotographs of meiotic stages were taicen with a Kam UBX 35 Him camera mounted on a .Reichert photomicroscope, with x40 objectives against a green filter. Drawings which are tracings from microphotographs were made of all the original pictures taken, these were indicated as figures* Due to technical problems developed by the photocopying machine used for this purposes, at the last stage of the xeroxing, tracings of the microphotographs of three species were made half the size of the microphotographs* These are: ncrosternum heegeri Fieb*, ■aeptus singularis (Dallas), and benia sp. A (Figs. 1-4, 9-12 and 4-3-46 respectively). In all the stages of meiosis examined, especially at the spermatogonial metaphase, the chromosomes were categorized into University of Ghana http://ugspace.ug.edu.gh different size groups, though in most cases absolute distinction between the different size groups was not usually possible* Species of Pentatomoidea examined, and their locality and date of collection are given in Table 1* 2n-numbers were determined for fifty-five (55) species of male Pentatomoidea, belonging to six families* Out of this number, microphotographs and tracings are presented for fifty-one (51) species. Distribution of 2n-numbers in the families of Pentatomoidea from literature ana those studied cytologically to date are presented separately in tabular ana in graphical form. Also distribution of 2n-numbers of families worked on in this study is similarly presented, specimens used in this work were identified at the ii^ ntomology museum of the ^oology apartment, University of mes to one pole University of Ghana http://ugspace.ug.edu.gh and the Y-chromosomes to the other pole. Chromosome complements were not determined for the females, so to distinguish the sex-chromosomes from the autosomes, and to establish the sex mechanism, the following criteria were used:- I. Diffuse stage of prophase I was studied and the sex chromosomes which are positively heteropycnotic at this stage were noted, unfortunately no good microphotograph of this stage was obtained. II. Metaphase II feature of autosomes forming a ring around sex- chromosomes provides an easy way of separating these two components of the karyotypes. III. Segregating patterns at anaphase II normally result in two types of chromosome distribution which gives an indication of tne mechanism of sex-determination: because while the autosomes divide equationally, the sex-chromosomes are segregated, and the Y-goes to one daughter cell, and the X- to the other daughter cell. University of Ghana http://ugspace.ug.edu.gh DBSCaaiFTIUN OF MEIOSIS University of Ghana http://ugspace.ug.edu.gh 3«3»1 Figs* 1-4: Acrosternum heegeri JTieb* : 1) Microphotograph# 2) Tracing (-J microphc) tograph size) of spermatogonial metaphase, showing diploid complements of 14 chromosomes, 3 long, 9 medium and 3 small elements. 3) Microphotograph, 4) Tracing (-J microphoto­ graph size) of Metaphase 1, arranged regularly on equatorial plate, smallest element is Y-chromosome. University of Ghana http://ugspace.ug.edu.gh •3*2 Figs, 5-8: Aeliomorpha spV griseoflava btal 5) Microphotograph, 6) 'Tracing of bpermatogonial metaphase, the chromosomes are mostly connected. Ik chromosomes (4 long, 7 medium and 3 small elements, 7) Microphotograph, 8) tracing of hetaphase II, polar view, autosomes arranged in a ring with sex chromosomes in a central position. University of Ghana http://ugspace.ug.edu.gh 3*3*3 figs. 9-12: Aeptus singular is (Dallas): 9) microphotograph, 10) tracing (■§• microphotograph size) of jJiakinesis, showing condensation of bivalents, 11) microphoto6raph, 12) tracing (-J microscope size) of Metaphase II, polar view chromosome size larger than that of iieliomorpha sp. (Pigs 7-8). University of Ghana http://ugspace.ug.edu.gh i? 18 17 JO /J 3*3-^ Figs* 13-18: Aethemenes chloris (westw.) 13) microphotograph, Ik) tracing of spermatogonial metaphase, showing Ik elements, 15) microphotograph, 16) tracing of Metaphase 1, with a sex chromosome in the centre and a ring of 6 autosomes around, 17) microphotograph, 18) tracing of Metaphase II polar view. Chromosomes smaller than in Metaphase I, with much condensation, and appear as rounded bodies. University of Ghana http://ugspace.ug.edu.gh c m • i 19 21 IOJU I V C 20 • * > /A 22 •5 Figs. 19-22: Aspavia acuminata (hont.) : 19) microphotograph, 20) tracing of spermatogonial metaphase chromosomes (3 long, 9 medium and 2 small, 21) microphotograph, 22) tracing of Jjiakinesis with 8 elements. Y-chromosome smallest among elements. University of Ghana http://ugspace.ug.edu.gh • £ * * \ * &7> 23 2k j [ 4 » 1 V 25 26 10/j 3.3-6 figs. 23-26: Aspavia armigera G?'abr.) 23) microphotograph, 2k) tracing of uiakinesis, 25) microphotograph, 26) tracing of Metaphase II, polar view, the ring-like arrangement of the autosomes on the first division metaphase plates regular, with a high degree of condensation and metaphase elements appear as very small and rounded bodies* University of Ghana http://ugspace.ug.edu.gh 27 28 v- ' - • - ^ Wr 51 % S 52 IO/J 3.3.12 Figs. 47-52: Carbula capito ^tal 47) microphotograph, 48) tracing of Diakinesis, 49) microphotograph, 50) tracing of hetaphase II, side view, sex chromosomes lying wide apart, 51) microphoto&raph, 52) tracing of Metaphase II, polar view showing sex chromosome pair. University of Ghana http://ugspace.ug.edu.gh 26 University of Ghana http://ugspace.ug.edu.gh 27 3*3*13 Figs. 53“58• Carbula carbula (Distant) 53) microphotograph, 54) tracing of i-dakinesis, elements larger than those of Carbular capito (Figs. 47-52), 55) microphograph, 56) tracing of Metaphase II, polar view, 57) microphotograph, 58) tracing of Metaphase II side view. Chromosome elements of Carbula carbula larger in size than those of jC. capito. University of Ghana http://ugspace.ug.edu.gh I op 3.3.14 Figs. 59-62: Carbula marginella bt§l 59) microphotograph, 60; tracing oT iiiakinesis, chromosome elements larger than those of 0. capito, but smaller than in 0. carbula, 61) microphotograph 62) tracing of Metaphase II, polar view, irregular ring-like arrange­ ment of 6 autosomes with sex chromosome pair in the central position. University of Ghana http://ugspace.ug.edu.gh j.3.15 figs. 63-66: Carbula melacantha total 63) microphotograph, Gk) tracing of topermatogonial metaphase v/ith Ik chromosomes (2 long, 9 medium and 3 small)* Chromosome elements elongated rather than rounded, 65) microphotograph, 66) tracing of Diakinesis. University of Ghana http://ugspace.ug.edu.gh loyu 3*3*16 Figs# 67-68: Carbula sp. nr. sjostedti 6chout. 67) microphotograph, 68) tracing of Metaphase II, polar view, autosomes in an irregular ring, sex chromosome in middle. Chromosome elements larger than those of Cm capito, smaller than C. carbular, and of about same size of C. marginella. University of Ghana http://ugspace.ug.edu.gh IOJIA 3*3«17 i^gs. 69-70 : Caura pugillator (Fabr.) 69) microphotograph, 70) tracing of Metaphase II, polar view, paired sex chromosome in central position of ring-like arrangement. University of Ghana http://ugspace.ug.edu.gh | „ I * . * * ^ 7 1 7 2 s / * 1 * - * * % < * 7 3 7 ^ ID J J 3*3*lS Figs# 71-74: Lriploxys bipunctata (iimyot et serville) 71) micropho- graph, 72) tracing of opermatogonial metaphase with 14 chromosomes (2 long, 9 medium and 3 small), 73) microphotograph, 7^ ) tracing of Diakinesis# University of Ghana http://ugspace.ug.edu.gh 3*3«19 Figs* 75-76; Amaaosana punctata Last* 75) microphotograph, 76) tracing of Spermatogonial metaphase with Ik chromosomes (2 long, 9 medium and 3 small). University of Ghana http://ugspace.ug.edu.gh 78 77) microphotograph, 78) tracing University of Ghana http://ugspace.ug.edu.gh 3.3*21 Figs. 79-82: i/ormia lutulenta total 79) microphotograph, 80) tracing of Metaphase II, polar view, 8l) microphotograph, 62) tracing of Late Anaphase II* University of Ghana http://ugspace.ug.edu.gh gpV |. % \ov 83 3.3*22 Figs. 83-84: Lfarmia sp. 83) microphotograph, 84) tracing of Spermatogonial metaphase v/ith 14 chromosomes (3 long, 9 medium and 2 small). University of Ghana http://ugspace.ug.edu.gh 84 85 3o3*23 Figs. 84-87• J-tymantis grisea Jen Haar: 84) microphotograph, 85) tracing of netaphase II, side view, 86) microphotograph, 87) tracing of Jiakinesis. University of Ghana http://ugspace.ug.edu.gh . s * • * V / t " 9^ 95 Figs. 88-95^ i^ysarcoris inconspicuus Ud*-b) 88) microphotograph, 89) tracmng of Diakinesis, 90) microphotograph, 91) tracing of wetaphase li, polar view, 92)microphotograph, 93) tracing of Metaphase II, side view with heteromorphic sex pair, 9^) micropho­ tograph, 95) tracing of i^ arly .anaphase. University of Ghana http://ugspace.ug.edu.gh 3.3*25 Figs. 96-97: Halyomorpha reflexa (sign) 96) microphotograph, 97) tracing of Metaphase II, side view, with heteromorphic sex pair. University of Ghana http://ugspace.ug.edu.gh Figs. 98-101: Lerida punctata (Palisot Beauvois) 98) microphotograph 99) tracing of Liakinesis, 100) microphotograph, 101) tracing of Metaphase II, polar view* University of Ghana http://ugspace.ug.edu.gh 3*3*27 Figs. 102-103: Macrorhaphis acuta (Dali) 102) microphotograph, 103) tracing of spermatogonia! metapha.se of Ik chromosomes. University of Ghana http://ugspace.ug.edu.gh 3-3*28 Figs. 104-109: Nezara viridula (.Linn.) 104) micropliotograph, 105) tracing of Diakinesis, 106) microphotograph, 10?) tracing of hetaphase I, sex chromosomes in central position ox irregular ring-like arrangement, 108) microphotograph, 109) tracing of ketaphase II, polar view. University of Ghana http://ugspace.ug.edu.gh r r w m p t . • ? : P 1 * 110 111 I k » 112 113 3*3*29 ■E'i&s* HO-II31 Nezara viridula var smaragdula (Fabr.) 110) micro- photograxDh, 111) tracing of Hetapiiase II, polar view. Chromosome elements smaller than in 1^. viridula, 112) microphotograph, 113) tracing of i*ietaphase II side view. University of Ghana http://ugspace.ug.edu.gh L________IOjU 3-3*30 *‘igs. 114-117: Nezara viridula var torquata (jmton) 114) microphoto­ graph, 115) tracing of hetaphase I, llo) microphotograph, 11?) tra­ cing of Hetaphase II side view, uenerally the chromosome elements are larger than those of JU. viridula and if. viridula var smaragdula. University of Ghana http://ugspace.ug.edu.gh 3.3#31 Figs. 118-123: Antestia sp. 118) microphotograph, 119) tracing of Spermatogonial metaphase of 14 chromosomes (2 long, 2 medium, 8 submedium and 2 small), 120) microphotograph, 121) tracing of Metaphase II polar view, 122) microphotograph, 123) tracing of Metaphase II, side view. University of Ghana http://ugspace.ug.edu.gh 3.3.32 Figs. 124-129: Antestia sp. s.l 124) microphotograph, 125) tracing of Pro-spermatogonial metaphase, 126) microphotograph, 12?) tracing of bpermatogonial metaphase with 14 chromosomes (2 long, 4 medium, 6 submedium and. 2 small), 128) microphotograph, 129) tracing of Diakinesis. University of Ghana http://ugspace.ug.edu.gh « • m m *i 131 9 -ki i 1J2 133 & \ * 135 3.3*33 Figs. 130-133: Antestia sp? immunda Linn. 130) microphotograph, 131) tracing of jjiakinesis, 132) microphotograph, 133) tracing of Metaphase I, irregular ring-like arrangement, 13 )^ micropho­ tograph, 135) tracing of Metaphase II siae view. I • # 134 University of Ghana http://ugspace.ug.edu.gh i' ^ i • \ • '1 ^ A 136 137 loy 3»3*3^ Figs. 136-1375 imtestiopsis sp. s.l 136) microphotograph, 137) tra­ cing of Diakinesis, 7 elements present with 1 missing probably due to imperfect squasning. University of Ghana http://ugspace.ug.edu.gh 0y *J • 0 1 • # # 138 139 \0/U 3*3*35 Pigs. 138-339i iriezodorus hybneri (Gmelin) 138) microphotograph, 139) tracing of Metaphase I. University of Ghana http://ugspace.ug.edu.gh 3*3*36 Figs. 140-143: ocotinophara fibulata (schouteden) 140) microphoto­ graph, 141) tracing of hetaxohase I, irregular chromosome arrangements, 142) microphotograph, 143) tracing of Hetaphase II polar view. University of Ghana http://ugspace.ug.edu.gh * / t m \ 4 144 145 \Ofd 3.3.37 i'igs. 144-145: bepontia misella (atSl) 144) microphotOarapn, 1^ 5) tracing of Diakinesis. University of Ghana http://ugspace.ug.edu.gh 3.308 Figs. 146-147: Tyoma verrucosa (Mont.) 1^ 6) raicrophotograph, 147) tracing of ketaphase II polar view. University of Ghana http://ugspace.ug.edu.gh 3*3*39 Figs. 148-14-9: Veterna sanguineirostris otal 148) microphotograph, 149) tracing of Metaphase II polar view; chromosome elements irre­ gular, arranged on equatorial plate. University of Ghana http://ugspace.ug.edu.gh 150 151 152 153 » * ' » 154 155 University of Ghana http://ugspace.ug.edu.gh 3«3*40 Figs. 150-1552 Macrina juvenca (Burm.) 150) microphotograph, 151) tracing of JJiakinesis, 152) microphotograph, 153) tracing of hetaphase I, 154) microphotograph, 155) tracing of Hetaphase II side view. University of Ghana http://ugspace.ug.edu.gh kl Figs* 156-159: Callidea duodecimpuntata otal 156) microphotograph, 15?) tracing of Diakinesis, Vjo) microphotograph, 159) tracing of Metaphase I. University of Ghana http://ugspace.ug.edu.gh 3*3*kZ Figs. 100-1632 Hotea subfasciata C^ estw.) 160) microjjhot ograph, 161) tracing of Hetaphase I, lb2) microphotograph, 163) tracing of lietaphase ii side view. University of Ghana http://ugspace.ug.edu.gh 3.3.43 Figs. 164-165; a-phaerocoris testudogrisea Stal 164) micropho- graph, 165) tracing of hetaphase I* University of Ghana http://ugspace.ug.edu.gh 3.3.44 Figs. 166-1712 frteganocerus multipunctata (de Geer) 166) microphotograph, 167) tracing of uiakinesis, 168) microphotograph, I69) tracing of hetaphase I, 170) microphotograph, 171) tracing of hetaphase II polar view* University of Ghana http://ugspace.ug.edu.gh 172 173 m i ** yw‘ j "•% r * % x Y % 174 175 ■y x 176 177 3.3 .45 i'ibs* 172-1772 Brachyplatys incertus (walker) 172) microphotograph, 173) tracing of Diakinesis, 174) microphotograph, 175) tracing of hetaphase I, 176) microphoto&raph, 177) tracin0 of Metaphase II, side view. University of Ghana http://ugspace.ug.edu.gh #. % 17B > - & % I * 179 10/J 3.3.46 Figs. 178-179: Brachyplatys testudoni^ra total 178) microphotograph, 179) tracing of Diakinesis. University of Ghana http://ugspace.ug.edu.gh 1j 1 * • * %! * , * > % V % Y 180 181 * 1 # v / . 182 183 1 or 3*3«^7 Figs. 180-183 ^Coptosoma nubila Germ. l80) microphotograph, l8l) tracing o£ opermatogonial metaphase with 12 chromosomes (2 long, 8 medium and 2 small), 182) microphotograph, 183) tra­ cing of Hetaphase II polar view. University of Ghana http://ugspace.ug.edu.gh 3«3*48 Figs* 184-185: Coptosoma stali hont. 184) microphotograph, 185) tracing of JJiakinesis. University of Ghana http://ugspace.ug.edu.gh * ** > % a H H R H 186 187 IDfJ 3.3.49 -Figs. l86-l87s Goridius cuprifer (tfestw.) 186) microphotograph* 187) tracing of i'letaphase II polar view. University of Ghana http://ugspace.ug.edu.gh * « • • • r * X ■BSWSKWBEfl 188 189 I0|U 3*3*50 Figs* 188-1892 Piezosternum calidum otal 188) microphotograph, I89) tracing of Hetaphase 1. University of Ghana http://ugspace.ug.edu.gh 190 191 I DM 3.J.51 Figs. 190-1915 iuacroscytus sp. 190) microphotograph, 191) tracing of bpermatogonial metaphase with 12 chromosomes (2 long, 8 medium and 2 small). University of Ghana http://ugspace.ug.edu.gh SECTION 4 DISCUSSION 4.1 SYSTEMATICS: The Pentatomoidea are essentially tropical or subtropical injects. They are mostly phytophagous although some are predacious. ’There is as yet no general agreement as on their higher classification. Some groups which are regarded as distinct families by some authors have been treated as subfamilies of Pentatomidae by others (e.g Schouteden, 19051 1910 and Kirkaldy, 1909). Recently, however with the results of work by investigators, such as Leston (1954a, 1954b, 1955)> Southwood (1956), bcudder (1959)1 Miyameto (I96I), Kumar(1962, 1965 , 1968a and b) and Cobben 1968, 1978) etc. the families formerly treated as subfamilies unaer Pentatomidae have been elevated to family status. Kirkaldy (1909) lists over 3,400 species of the Pentatomidae and includes them under subfamilies. Brues, Helander and Capenter (1954) erect a superfamily the Scutelleroidea/Pentatomoidea, with the following families under it: Cydnidae, Corimelaenidae, Plataspidae, Scutelleridae, Podopidae, and Pentatomidae. China ana niller (1959) raised the Aphylidae and Phloeidae to family status, thus further reducing the subfamilies under Pentatomidae as listed by Kirkaldy (1909)* Leston (1958) represented the central group of Pentatomoidea families with the series: Tessaratomidae, Scutelleridae, Cydnidae, Aphylidae, University of Ghana http://ugspace.ug.edu.gh □0 a < oca;in A Q. a JS F R E Q U E N C Y (/) o-, ^ L, j> o 3 o a U r o s t y l i d a e ■ ■ i E u m e n o t i d a e j D i n i d o r i d a e B r a c h y p l a t i d a e S c u t e l l e r i d a e ! A c a n t h o s o m a t i d a e C y d n i d a e T e s s a r a t o m i d a e P e n t a t o m i d a e F A M I L Y CD _* Xs H o T 12 N> K ) 1 2 31 l hO r o r-o — ‘ * - j c n cr> c n 4^ r o O C T ) 2 N - N o T | “ - CD 1 ^ — © s OJ —* i s ~ * C 7 ) - » N ) ^ 3 0m o K E Y University of Ghana http://ugspace.ug.edu.gh Urostylidae Eumenotidae Dinidoridae Brachyplatidae Scutelleridae >OPD Or 8 S . ° 1 Cydnidae Tessaratomidae Pentatomidae FAM 1 LY K>—a —* —V —» —* —a CO —* —> —»NJ N) —k —k —» —* —k zCD 4N b^- K) hO K) —* K> NJ '-0 izO - — —Ao CD CD — - U) oo FREQ. KEY University of Ghana http://ugspace.ug.edu.gh Fig.19^ Distribution of 2N~ Numbers in 170 species of Pentatomidae Cytologically studied and already published. K> FREQUENCY CT) CD O K> ^ C.O OJ O N> J ______! i : c j K>cn ro00 : = r Podopinae 1 Amyoteinae Pentatominae . ........................... cnczCD > r~*< hO--—4> n ) ro ZCD rsj *--3(T)CJ>U 14NrOO Cn jIZ • 0—V *~n—f. OJ ZO—* u> 0 —»■—' cn —4 — * —^ 1 ~s> r nO m -< University of Ghana http://ugspace.ug.edu.gh JTig.195 Dfsiribution of 2N-Numbers in 211 species of Pentatomidae Cytolog ically studied up to date . z c CD m hO FREQUENCY u i — 4 •* ”*0~) OO O KJ> CD rx) —» -* ”11X)mo 7Km *< University of Ghana http://ugspace.ug.edu.gh Plataspidae, Pentatomidae, Dinidoridae, Urostylidae and Acanthosomatidae, with Phloeidae as an early off shoot of Tessaratomidae. Recently, the book entitled, "The Insects of Australia'1 (Mackerras, 1970) includes the following families occurring in Australia under the super­ family Pentatomoidea: Tessaratomidae, i)inidoridae, ocutelleridae, Plataspidae, Aphylidae, Lestoniidae, Urostylidae, Cydniaae, acanthosomatidae, s/ and Pentatomidae. A new family, Thaumastellidae btys has recently been erected by £tys (1964) on account ox its primitive chorionic structure, as revealed by the eggs of Thausmastella aradoides. otysf(l964) conclusion that Thausmastellidae must be regarded as an early offshoot of the pentatomoid stock is confirmed by Cobben (1968). 4.2 KARYOTYPE aIM'jj CLARIFICATION: From the literature, the chromosome number of only 217 species of Pentatomoidea is known (Fig. 192, appendix Tables 3» 6-11). There may be a few errors of omission which may be attributed to the limited library facilities available to me. The present work has examined forty-nine (49) species where chromosome number had never been worked out earlier. In all fifty-five (55) species belonging to six families of Pentatomoidea were examined, out of this only six species had previously been examined. 'This brings the number of species of Pentatomoidea with known chromosome number to 266 (Fig. 195)* 4.3 family (Fi^s. 1 - 155): 'This is the largest family among the group Pentatomoidea, and probably presents the most common species among its members. Until University of Ghana http://ugspace.ug.edu.gh recently almost all the other families of the Pentatomoidea we re placed under the Pentatomidae as subfamilies, but following morpho— taxonomical studies on genitalia, alimentary canal, salivary glands, and wing venation, etc* the Cydnidae, 'i’essaratomidae, LdLnidoridae, Brachyplatidae, Scutelleridae, Acanthosomatidae and Urostylidae, were all raised to family status. The Pentatomid bugs, being easily available, have had their cytological data relatively well extended* However, since the review of Manna (1958), little work seems to have been published on their chromosomes* !Ehe Pentatomidae is more or less uniform with regard to chromosome number with the elevation of the subfamilies Brachyplatinae, Cydninae, Tessaratominae, ocutellerinae, and i-dnidorinae raised to family status* Uhis family is characterised by Ik chromosomes (modal number), XX:XY type of sex determination, absence of m chromosomes in the species so far studied, a characteristic disposition of the first and second metaphase chromosomes, in which the autosomes are arranged in the form of a ring around the periphery of the spindle, and the sex-chromosomes usually lie at the centre (Manna, 1958). Two hundred and eleven species (211) of four subfamilies of Pentatomidae are so far known cytologically (Appendix i'able **)• She Amyoteinae (Asopinae) are known cytologically through the study of fifteen species (i?'ig* 195)* In tne males, three diploid numbers of University of Ghana http://ugspace.ug.edu.gh 12, 14 and 16 have been reported, of which 14 seems to be the original (Manna, 1951, 1958, Makino 1951)- Manna (1958) explains this by the fact that most of the species so far studied (11 out of 15) possess this number, one species Uechalia patruelis has 12 chromosomes. '.Che number 16 is found in three species of ivpateticus (Podisus). Manna (1958) suggests that since two species of Apateticus and one of Oechalia have 14 chromosomes, it can be concluded that the 16 chromosome species in the latter are derived from the basic 14 in Amyoteinae. A species each of Podopinae and Phyllocephalinae (from this study) is known, and in each case the chromosome number is 14. Pentatominae possess the most diversified chromosome numbers, ranging from six to twenty-seven among the one hundrea and ninety- four (19^ ) species studied up to date (appendix 1‘able 4, Jfig. 195)* As Manna(1958) puts it “it is apparent from the cytological data that this subfamily includes heterogenous groups’1 • whilst all the species in the tribes of Jr'entatominae, namely, j-dscocephaiini, iiidessini, Halyini and ociocorini, examined have 14 chromosomes (Hanna, 1958). The tribe Pentatomini evinces a marked conservatism as to changes in the number and benaviour of its chromosomes. Of the one hundred and minety-four (194) species examined, one hunared and seventy (170) have chromosome number of 14, eleven possess 16, in six the complement is 2b, in three the number is 12 each, and one University of Ghana http://ugspace.ug.edu.gh >- o 2 UJZD O LU DC Ll 10 8 H 6 - U - 2 t— r— i— r 10 12 KEY SUB FAMILY 2N-No FREQ- Scu teller inae 1 2 7 Pachycorinae 12 1 Eurygastrinae 12 7 2N-NUMBER Fig.196 Distr ibution of 2N -N umbe rs in 15 species of Scute l ler idae Cyto log ica l ly s tud ied and a l ready pub l ished . 1 0 * 8 ->■ 0z 6 - Ll )D A -O Ll) DC 2 -Li- r r n 2N-NUMBER KEY ,l|i SUB FAMILY 2N-No FREQ- Scutellcnnae 12 10 ! Pachycorinae 12 1 I % Eurygastrinae 12 7 12 1 114 iTi^ .197 / D is t r ibu t ion of 2N -N um be rs of Scutelleridae Cyto log ica l ly s tud ied to da te . University of Ghana http://ugspace.ug.edu.gh species each contains 6, 10, 15 «md 27 chromosomes (i?ig. 195)• &o far as our evidence goeSf the few deviations from the characteristic chromosome of l*f, may be due primarily to processes^" involving fusion and fragmentation (bchrader, 19 7^ » Schrader and Hughes-bchrader, 1956)* In all Pentatomini (except Thyanta calceata) the males have a XY sex determining mechanism and in all it is in the second meiotic division that the a and Y segregate from each other and enter separate spermatius. It should be mentioned here that the ways in which the cytological changes here described have come about are still a matter of conjecture (Hughes-bchrader and Schrader, 1956). There is nearly always a rather even gradation in size from the largest to the smallest chromosomes in a complement. This suggests that extensive rearrangements have not been common in the chromosomal evolution of the Pentatomini. Since species with less than 14 chromosomes are even rarer, it would appear that factors additional to fragmentation and fusion are at work to make such changes permanent, and that these are not often present tochrader, 19 -^7)* FAHIliY SCUTfiLT»< .RIjjah* (iTigs. 196 and 197): There is considerable disagreement over the status of this group. Heuter (1912), van iXizee (1917) * bcudder (1959) * Gobben (1968) and several others assign it a family status, while others like iu.rkaldy (1909)i Pruthi (1925), Southwood (1956) and liiyamoto (1961) have considered it to be of subfamily rank, bouthwood (1956), in dealing University of Ghana http://ugspace.ug.edu.gh w i th t h e e g g s o f P e n t a to m o id e a , t r e a t e d t h i s g ro u p a s a s u b f a m i l y , b u t C obben (1968) p o i n t s o u t t h a t ’’s c u t e l l e r i d e g g s , c a n b e d i s ­ t i n g u i s h e d f ro m P e n ta to m id e g g s b y t h e p e r s i s t e n c e o f som e l o n g i n t e r n a l m i c r o p y l a r c a n a l ” a n d t h a t ‘‘c l e a r f a m i l i a l d ic h o to m y i s s u g g e s t e d s i n c e som e P e n ta to m id s p e c i e s h a v e t h e sam e lo w a n a g e n e t i c l e v e l f o r t h e i r e g g s t r u c t u r e s ( a r r a n g e m e n t o f m i c r o p y l e s a n d in c o m p le te p s e u d o p e r c u lu m ) a s s c u t e l l e r i d a e b u t l a c k t h e i n t e r n a l c a n a l* ', t h u s t h e s c u t e l l e r i d a e c a n n o t b e p l a c e d u n d e r t h e P e n t a t o m id a e . Kumar (1962, 1964, 1965), w o rk in g on t h e g e n i t a l i a a n d s a l i v a r y g l a n d s o f t h i s g ro u p a n d r e v i e w in g o t h e r e v id e n c e c o n f i rm e d t h a t t h e s c u t e l l e r i d a e , i n d e e d d e s e r v e s a f a m i l y s t a t u s w i t h i n t h e s u p e r f a m i l y P e n t a to m o id e a . Chrom osom e n u m b er o f e i g h t e e n s p e c i e s , b e l o n g in g t o e l e v e n g e n e r a , i n c l u d i n g t h e t h r e e s p e c i e s , C a l l i d e a d u o d e c im p u n c ta t a ( s t a l ) , H o te a s u b f a s c i a t a (W estw ) a n d a t e g a n o c a r u s m u l t i p u n c t a t a (d e G e e r ) e x a m in e d i n t h i s s t u d y ( F i g s . 156 - 171) i s k now n . A l l t h e s p e c i e s p o s s e s s 12 ch rom osom es ( F i g s 196-197)* w h ic h c a n t h e r e f o r e b e t a k e n a s t h e c h a r a c t e r i s t i c n u m b e r f o r t h e S c u t e l l e r i d a e . n o w e v e r , a c c o r d i n g t o M anna (1958), o f t h e f o u r o r f i v e s p e c i e s o f M i r y g a s t e r , m a u ru s i s r e p o r t e d b y G e i t l e r (1938) t o h a v e i n d i v i d u a l s w i th 14 ch ro m o so m es (A p p e n d ix T a b le 8 ) . A l l t h e s p e c i e s o f t h e s c u t e l l e r i d a e a r e s a i d t o h a v e t h e sam e g e n e r a l p l a n o f m e io s i s w i th a k Y ik X t y p e o f s e x d e t e r m in a ­ t i o n c o m p a ra b le w i th t n a t o f t h e p e n t a to m id b u ^ s (ivia n n a , 1958). University of Ghana http://ugspace.ug.edu.gh 4*5 FAMILY BRaCHYPLATI-UAE (PLATASPIDAfl) S It is usual to give it a family status these days. From morpho- taxonomical studies on the genitalia and salivary glands, Kumar (1962), observed that the Brachyplatidae are a specialized group exhibiting close resemblances to Cydnidae, but the differences in the two are sufficient enough to create different families for each of the two. Recent authors like Brues, Melander and Gapenter (195^)t Manna (1958), China and Miller (1959)» Mackerras (1970) in the “Insects of Australia1*, and Cobben (1978), all make reference to the Plataspidae. The cytological knowledge of ten species is known, four of Brachyplatys and six of Coptosoma. This includes the two species Brachyplatys incertus (walker) and Coptosoma nubila Uerm where chromosome numbers have been added on from this study (Figs. 172-185)• 'The Brachyplatidae is characterized by having 12 chromosomes (Fig. 192) with the usual ^Y:jQ£ sex determination. 'The behaviour of the sex chromosomes during meiosis follows the usual pentatomid pattern. Manna (1958) reports that the meiotic cells are smaller than those of pentatomid bugs, however this study found them to be about the same, except in Coptosoma etali (Mont), where the cells appear larger. The significance of these observations would perhaps be better appreciated wiien lot more work been done on this family. It is suggested that more serious thought be given to devising a project on the Plataspiaae alone, so that more information on their chromosomes would be known for more substantive University of Ghana http://ugspace.ug.edu.gh c y t o l o g i c a l d e d u c t i o n s t o be made on t h e s y s t e m a t i c s t a t u s o f t h i s f a m i l y . 4.6 DlMlDQRl^^ S in c e 1© 70 , t h e D i n i d o r i d a e h a s b e e n c o n s i d e r e d a d i s t i n c t g ro u p p o s s e s s i n g c e r t a i n s p e c i a l f e a t u r e s ( S i n n a d u r a i , 1979)* Lethierry et Severin (1893)t in their catalogue of Hemiptera classified it as a ’’subfamily Dinidorinae". distant (l88l), called it uinidorinae and the group was classified under this name until the middle of this century. More recent authors, such as Leston (1955)$ Pendergrast (1957)* Scudder (1959)♦ Miyamoto (1961), Kumar (1962, 1965)? Cobben (1968, 19?8) etc. have on a variety of grounas accorded this group a family status. 'Ih e f a m i l y D i n i d o r i d a e i s now d i v i d e d i n t o tv/o s u b f a m i l i e s , t h e i ^ i n id o r i n a e a n d t h e M eg ym inae , e a c h o f w h ic h i s f u r t h e r s u b d i v i d e d i n t o tw o t r i b e s ( b i n n a d u r a i , 1 9 79 )* i h e D i n i d o r i n a e i n c l u d e s D i n i d o r i n i a n d ‘f h a l m i n i , a n d t h e M egym inae h a s M egym in i a n d j^ u m e n o t in i . C o r i d i u s , t h e l a r g e s t g e n u s w i t h 59 s p e c i e s a n a i d n i d o r come u n d e r D i n i d o r i n i . The common g e n u s u n d e r M eg ym inae , iviegymenum i s a l s o p l a c e d u n d e r t h e t r i b e M egym in i w h i l e .b u m e n o t in i i s r e p r e s e n t e d by t h e m o n o ty p ic g e n u s M -uneno tes . C o r i d i u s se em s t o b e t h e m o s t a d v a n c e d a n d h i g h l y s u c c e s s f u l g r o u p . A c c o rd in g t o S i n n a d u r a i (1979)* t h e S u b f a m i ly M eg y im in ae i s s u p p o s e d t o show p r i m i t i v e c h a r a c t e r s i n e x t e r n a l f e a t u r e s s u c h a s t h e a n t e o c u l a r p r o c e s s e s , p r o n o t a l t u b e r o s i t i e s a n a t n e l o b e d o r University of Ghana http://ugspace.ug.edu.gh tubercled connexiva. The male genitalia has a poorly developed conducting chamber and a single pair of conjunctival processes v/hich are quite elongated in the genus hegymenunu The sack-like spermatheca seen in this genus is more primitive than the genus Jiiumenotes v/hich has less elaborate external features# as has rightly been pointed out by Sinnadurai (1979)> the evidence for the affinities of the uinidoridae to other families of the Pentatomoidea have been drawn mostly from work carried out on the single species Goridius ianus or the genus Goridius and occassionally on the genus hegymenum. Close similarities to the family Pentatomidae have been shown by the work of several authors. The trichobothrial arrangement in the abdomen of J-anidoridae is similar to Pentatomidae (Schaeffer, 1966). Work on the chromosomes by Heston (1955) leads to a similar conclusion. However, according to liumar (1962} the genitalia and salivary glands of Ldnidoridae differ considerably from the Pentatomidae. There is also to some lesser extent some similarity of Dinidoridae to Tessaratomidae, ocutelleriaae and Acantliosomatidae (binnadurai, 1979)* Cobben (1968) suggests that such overlap means no mutual relationship but parallelism. -Sinnadurai concludes her treatise by saying that the Dinidoridae is probably a family that differentiated from the Pentatomid stock at an early stage and became a specialised group. Cytologically, only seven species (including the one of this study, Coridius cujorifer (Figs. 186 - 187), belonging to three genera, Goridius (Asjpongopus), idnidor and hegymenum have been studied. These University of Ghana http://ugspace.ug.edu.gh t h r e e g e n e r a h a v e d i f f e r e n t d ip l o id , n u m b e rs (F ig # 19 2 ) , 1 4 i n f o u r s p e c i e s o f C o r i d i u s , 2 0 i n tw o s p e c i e s o f Hegymenum a n d 2 1 i n D in id o r r u f o c i n c t u s . Of t h e s e t h r e e g e n e r a , tw o o f th e m ( C o r i d i u s a n d D in i d o r ) b e lo n g t o t h e s u b f a m i l y , D i n i d o r i n a e , w h i l s t Megymenum i s i n t h e s u b f a m i ly h e g y m in a e c r e a t e d b y b i n n a d u r a i (1979)* The f a c t t h a t t h e tw o s p e c i e s o f Megymenum e x a m in e d h a v e t h e sam e ch rom osom e num ber o f 2 0 , a n d t h e f o u r s p e c i e s o f C o r i d i u s a l s o p o s s e s s a ch rom osom e num ber o f 1 4 , s e em s t o s u p p o r t t h e c r e a t i o n o f t h e s e c o n d s u b f a m i ly * H o w ev e r, t h e r e i s t h e q u e s t i o n a s t o why t h e D in i d o r a l s o o f t h e sam e s u b f a m i l y a n d t r i b e a s t h e C o r i d i u s h a s a d i f f e r e n t ch rom osom e n um ber o f 2 1 , a n d n o t I k a s C o r i d i u s ? D oes i t s u g g e s t s t h e c r e a t i o n o f a n o th e r s u b f a m i ly ? I t i s t o b e n o t e d h o w e v e r t h a t D . r u f o c i n c t u s i s o f d i f f e r e n t g e n u s th o u g h o f t h e sam e s u b f a m i l y a n d t r i b e a s O o r id iu s « F u r t h e r a c c o r d i n g t o S c h r a d e r ( 19 ^ 7 ) ,£♦ r u f o c i n c t u s i s s u p p o s e d t o h a v e a m u l t i p l e s e x ch rom osom e m e c h a n ism , h e n c e i t s h o u l d n o t b e s t r a n g e f o r i t CD. r u f o c i n c t u s ) t o e x h i b i t a d i f f e r e n t ch rom osom e num ber* b in c e i t i s p o s s i b l e f o r on e t r i b e o f a s u b f a m i l y t o p o s s e s s d i v e r s i f i e d chrom osom e n u m b e r s , a s i s fo u n d i n t h e P e n t a t o m in i o f P e n t a t o m in a e . ’The d i f f e r e n c e s i n chrom om osom e n u m b e rs o f hegym enum a n d G o r i d i u s c o n f i r m t h e m o rp h o - ta x o n o m ic a l s t u d i e s by b i n n a d u r a i ( 1 9 7 9 ) , w h ic h l e d h e r t o c r e a t e tw o s u b f a m i l i e s , D i n i d o r i d a e a n d w e g y in i n a e . The p r e s e n t c y t o l o g i c a l d a t a a r e h o w e v e r r a t h e r m e a g r e . I t i s h o p e d t h a t f u t u r e w o rk w i l l c o n c e n t r a t e m o re on t h e ■ L'inidoriciae, t o p r o v i d e a d e q u a t e c y t o l o g i c a l i n t o r m a t i o n t o s u p p le m e n t t h e m o rp h o - ta x o n o m ic a l University of Ghana http://ugspace.ug.edu.gh 4 -i>-o 3 -2UJD 2 -C3 1 -UJccLl. KEY SUB FAMILY 2N-No FREQ Tessa ratom inae 12 2 Oncomerinae 1 4 1 2 11 12 14 2N-NUMBER Figk'198D is t r ib u t io n of 2N-Number in 4 species of Tessaratomidae Cyto logica l ly s tud ied to d a te . 8 -I >o 6 -zLU 4 -Z>O 2-q:Ll i— r 10 12 14 KEY SUB FAMILY 2N-Np FREQ Oncomerinae 1 2 1 Tessaratominae 1 2 2 2N-NUMBERF ig .199 Dis tr ibu t ion of 2N -Numbe rs in 3 species of Tessaratomidae University of Ghana http://ugspace.ug.edu.gh studies of Sinnadurai (1979) £or the establishment and creation of the two subfamilies# 4.7 Family T^SARaTOBIjjAL (Figs 198 and 199) The status of this group within the Pentatomoidea is undecided# Leston (1954a) and Lupius (1949) bave raised it to family status, with three constituent subfamilies: the Oncomerinae, Natalieolinae and the Tessaratominae. In Leston*s (1955) opinion, one of its genera Piezosternum, displays certain features, r,very suggestive11 of corefiid affinities. Kumar (1968b) however, observed that the Oncomerinae shows more similarities to Tessaratomini than to any other group of Pentatomor- pha and is thus placed close to Tessaratomini in the Tessaratomidae. He further observed that the uncomerinae is a homogenous group and bears no relationship whatsoever to Coreoidea, noting that similarities between Tessaratomidae and Coreoidea, emphasised by certain authors are superficial and based on an inadequate investigation of far too few representatives of the groups concerned. Cytologically, very little work has been done on this family. Until this study, only three species were known cytologically with a chromosome number of 12 (Appendix Tables 3 ) . 'The species are saevus (otal), lessaratoma javanica (Thunberg), and Piezosternum subulatum (otal). The species examined in this study, Piezosternum £alA,4vuift t has a chromosome number of 14 (Figs 188-189) with an XY:aA type of sex determination. If the basic chromosome number University of Ghana http://ugspace.ug.edu.gh of the Tessaratomidae is 12, then they can be placed in the same category as the Plataspidae, scutelleridae and Cydnidae etc. and may thus have some similarity of these, however the 14 chromosome number of Piezosternum places it close to the Pentatomidae. The present data, however, is too scanty to be able to predict any basic chromosome number of this group. A cytological investigation of the rich Australian Uncomerine fauna is especially desirable. 4.8 Fa h ILY CYlM I j Ahj The family now consists of oehirinae, Cydninae and Coriomelaeinae. huraar (1962) suggested that Cyuninae and Coriomelaeinae should be raised to family level, having treated oehirinae as one of the Cydninae. The Cydnidae is observed to possess three pairs of conjunctival processes# and in this respect, show affinities to Scutelleridae, J-dniaoridae, and Tessaratomidae (Leston, 195^a)« Male reproductive organs however tend to indicate its affinities to Plataspidae (Kumar, I962), while ocudder (1959) on ovipositor of Coriomelaeinae observed its affinities to certain Tessaratomidae and Pentatomidae. However, both Scudder (1959) and Pendergrast (1957) noted that Thyreocoris does not show affinities to Cydninae, which clearly indicates that Thyreocoris is placed in a wrong subfamily as suggested by ocudder (1959). We have cytological knowledge of eight species, seven have chromosome number of 12, and one stibaropus molginus has Jl. whilst those with the 12 chromosomes number have aY;^ type of sex mechanism. University of Ghana http://ugspace.ug.edu.gh molginus, has X^J^Y sex chromosomes in the males (appendix Table ?). The only species, hacroscytus sp. worked on in this study has a chromosome complement of 12 (Pigs 190-191) • basic chromosome number of Oydnidae is 12, and this places the family closer to Plataspidae and Tessaratomidae, as the morpho-taxonomical stuaies of Lest on (1954b) and Kumar (1962) have already revealed and this fact is further confirmed by the available cytological information. 4.9 O g M iftmiiilHiS UP PiLNTATul'iuI.UkA Three other families in literature where cytological information is known, but specimens of which could not be examined in this study, because of unavailability are: Acanthosomatidae (Appendix Table 6), •bumenotidae and Urostylidae. bix species belong to three genera of Acanthosomatidae namely, Acanthosoma, Plasmostethyus and ilasmucha have been examined cytologically, with chromosome number of 12, some individuals of i£Lasmucha are however said to have 18 chromosomes (Parshad, 1957a). Manna (1958) observes that Acanthosomatidae are distinct cytologically from the Pentatomidae, and are characterised by 12 chromosomes. On the basis of this, the Acanthosomatidae can be said to show affinities to the other groups also with diploid members of 12, like the Brachyplatidae and the Scutelleridae. This is in agreement with Leston (1958), who placed these groups close together with the Tessaratomatidae. Kumar®s (I962) observation from studies of salivary glands also suggests that the Acanthosomatidae may have affinities with bcutelleridae. borne authors (Pendergrast, 1957; University of Ghana http://ugspace.ug.edu.gh Kumar, 1962) stressed that Urostylidae, iicanthosomatidae and dinidoridae have several genital features in common with Pyrrhocoridae, but Cobben (1968) thinks such overlap means no mutual relation, but merely parallelism. Lumenotidae until now based on a single genus Lumenotes is accorded a tribal status by binnadurai (1979)* The other family Urostylidae is known cytologically by only one species Urostylis pallida of Urostylidae has 16 chromosomes. In a study based on diploid numbers, Leston (1958) kept Urostylidae as family perhaps related to his 2n = 12 group (Tessaratomidae, Scutelleridae, etc). Until Parshad (1957)* found that 2n = 16 in Urostylidae, Leston took this family off his more primitive pentatomoid stock at a point later tnan differentiation of the iicanthosomatidae. According to Kumar (1971)1 Urostylidae seem to have differentiated at a much earlier stage. It appears that Urostylidae and Pyrrhocoriclae diverged from a Pentatomoid-Pyrrhocoroid stock at an early stage in the evolution of Pentatomomorpha, possibly close to a point where iuradoidea branched off. As Kumar (1971) admitted, the above explanation is not fully satisfactory, however in the groups discussed above, there are common features which are the result of parallel evolution in response to functional demands. 'There is far more room for further work cytologically to provide supplementary ideas on affinities of most of the groups here discussed. From the preceding discussions of individual families, the following karyotype groups appear to emerge, namely: Ik and 12. Uncter University of Ghana http://ugspace.ug.edu.gh the group 14 is the family Pentatomidae, whilst the families Scutelleridae, Cydnidae and Plataspidae fall under the karyotype group 12. 'These agree with results of raorpho-taxonomical studies of workers such as Southwood (1956)* Pendergrast (1957)» Scudder (1959)i Miyamoto (1961), Kumar (1962, 1964, 1965* 1968 etc) and Cobben (1968, 1978) that these later groups show affinities to each other. No basic chromosome number can however be cited for the other families of Pentatomoidea, that is JJinidoridae, Tessaratomidae, Acanthosomatidae and Urostylidae because the cytological data for each of these are too scanty, for any sound conclusions to be arrived at. Generally, however, it appears that the Pentatomoidea is characterized XY:XX sex deter­ mining mechanism. The males invariably possess a single Y-chromosome, but species with more than one X-chromosome are not uncommon. It has been suggested that the 12+aY chromosome mechanisms, may be regarded as representing the ancestral pentatomoid stock, perhaps before its divergence into the various subfamilies and families (Manna, 19585 Leston, 1958; Parshad, 1957b). However whilst both Hanna (1958), and Leston (1958) put 2n = 14 as the point of origin, .Leston considers 12 as the modal number of origin. Generally, from the results ana discussions, it is found out that the cytological information gathered from this work was in agreement with most of the results of morpho-taxonomical studies in determining the affinities of the families of the Pentatomoidea to one another. University of Ghana http://ugspace.ug.edu.gh There were no obvious contradictions of the cytological data presented and information obtained from the results of work already known from morphological studies. Lven at places where the cytological information was too little and dispersed, there were indications that with further work more light could be thrown on the problems considered in this discussions. University of Ghana http://ugspace.ug.edu.gh Fig.200 Distribution of 2N-Numbers in the 6 families of Tropical Penta- tomoidea worked on in this study. f r e o u e n c v hoz d 2 ca m XJ ro CD O r o O ishO 4S CD C ydnidae . . .. . w onI T ■o 0 a . Of t D inidoridae Tessaratom idae Scutelleridae ............... -......... .... P entatom id ae FAM ILY ........ hO z—A —» —k __k —» —* 1r o t o NJ - y O "T I 4> 1 0—ft 4N —- -IN 4>- mo IXm -< University of Ghana http://ugspace.ug.edu.gh SECTION 5 SUMMARY 55 species of male Ghanaian Pentatomoidea belonging to six families were cytologically investigated, 49 of them being reported for the first time, i'he spermatogonial chromosome numbers (2n), and sex determining mechanisms of all the species were elucidated. Microphotographs and tracings of karyotypes observed for each species were presented as figures (1 - 191)* Histograms of 2n- numbers of the various families were also constructed. Information gathered was pooled with the information in literature, analysed and discussed. In all 21? species of Pentatomoidea are known cytologically from the literature. I’he present work has increased this number to 266 species. Course of meiosis, was found to be typically heteropteran in all the species examined, The family Pentatomidae was found to exhibit diploid numbers ranging from 6-27 with 14 as the modal number. It has an aY:XX type of sex determining mechanism. 211 species of Pentatomidae belonging to 4 subfamilies are so far known, 41 of these being contribution from this study. The Scutelleridae and Plataspidae are cnaracterised by 12 chromosomes, with an aY:aa type of sex determining mechanism, and a general plan of meiosis comparable with that of the Pentatomid bugs. 18 species of scutelleridae including the three species, Callidea duodecim University of Ghana http://ugspace.ug.edu.gh punctata (Stal), Hotea subfasciata (Westwood), and Steganocerus multipunctata (de Geer) were examined in this study, and 10 species of Plataspidae also including the two species, Brachyplatys incertus (Walker), and Coptosoma nubila Germ from this study are known cytologically. 6. Ihe basic chromosome number of 12 for both the Plataspidae and the Scutelleridae, were found to confirm results of morpho- taxonomical studies which suggests some affinities betv/een the two. 7« 'The Dinidoridae is known cytologically from only seven species with chromosome number of 14, 20 and 21. The difference in chromosome numbers of Coridius (14), Dinidor (21) and Megymenum (20), seem to support the creation of a new subfamily Hegyiminae, by Sinnadurai (1979)* dinidor rufocinctus is said to have a multix^ le sex chromosome mechanism, -kumenotis obscura now placed under the newly created subfamily Wegyiminae by Sinnadurai (1979) Has a chromosome number of 14 and a XY:aa sex determining mechanism. 8* 4 species of Tessaratomidae are known cytologically with a chromosome number of 12 for three species and 14 for one species, Piezosternum calidum (bt£l) examined in this study. ‘The later has a XY:XX sex determining mechanism. 9* 8 species of Cydnidae including the lone one of this study (hacroscytug sp) are known cytologically with a diploid complement of 12 for seven species, and one species otibaropus molgimus (ochiodte) has a complement of 31. It has a aY:Xa sex determining mechanism. University of Ghana http://ugspace.ug.edu.gh 10. ihe Acanthosomatidae is characterized by 2n=12 chromosomes with a XYsaX sex determining mechanism. £>ix species are known cytologically belonging to three genera: Acanthosoma, iiilasmucha and the Llasmostethus. 'i*he possession of 12 chromosomes as a basic number may indicate itsaffinity to the Plataspidae, ocutelleridae and probably Cydnidae, as morpho- taxonomical studies have suggested. 11. Urostylidae which was considered by Leston (1 9 5 8 ) as a family related perhaps to his 2n=12 group (Tessaratomiciae, £cutelleridae etc) is known to have 1$ chromosomes by i'arshad (1 9 5 7 a) in Urostylis pallida. More work is therefore needed to clarify the correct affinities of Urostylidae. 12. ‘Ihere is still more room for lot more work cytologically to provide more light on ideas on affinities of Pentatomoid families and sub­ families. University of Ghana http://ugspace.ug.edu.gh SECTION 6 LITi^ ATUKH CITiiiD AKINGBuHUNGBL, a .£. 1974. Chromosome numbers of some North American Mirids (Heteroptera :lviirictae) Can. J. Genet, and Cytol. 16: 251-256• SaNERJ-i^ , M.K. 1958. A study of the chromosomes during meiosis in twenty-eight species of Hemiptera (Heteroptera, HomopteraJ. JProc. o^ol. qoc. Calcutta. II: 9*31• BHUjiiS, C.T.f , A.L and CaHP^T j^:, F.i-i. 1954* Classification of Insects, bulletin of the auseum Comparitive zoology. Vol. 108. p.l80. t^ AKAYU'J, J. 1950* Nombres et aes ovarioles dan les ovares des Hemipteres-Heteropteres. bull, aus. hist, nat. Paris (2). 22: 470-475* CHINA, rt.iii. 1955. A reconsideration of the systematic position of the family Joppeicidae Keuter (Hemiptera: Heteroptera) with notes on the phytogeny of the suborder. Ann. Mag* flat. Hist. (12) 8 :^ 353-70* CHINA, w.iii and hLLucjxt, N.C.j^. 1959* Check-list and keys to the families and subfamilies of tne Hemiptera-Heteroptera. Bull. Brit. ms U'Jat. Hist.) entomology 8^: 1-45. COBujiiM, H.H. 1968. Evolutionary trends in Heteroptera t^. I ■^ gg6* architecture of the shell, gross embryology and eclosion. Cen r^e Agri. iAxbl. i^ oc. waRenin.^ en 475 University of Ghana http://ugspace.ug.edu.gh 8. CQBBiilM, K.H. 1978. evolutionary trends in rieteroptera Ft II. Mouthparts-structure and feeding strategies. Centre agri. Publ. Doc. n/ageningen. 407 pp* 9* DIoTaijT, w.L. 1881. Insecta jtfhynchota Hemiptera-Heteroptera. Biol, centr. i-uner. het. 1: 1-462. 10. DUPIUb, C. 1949* Les Asopinae de la faune framcais (Hemiptera Pentatomidae) Lssai sommaire de synthese morphologique, systematique et biologique. Kev. Franc. ent. 16: 233-250. 11. jiiVAkb, J.W. 1946. A natural classification of leaf-hoppers (Jassoidea, Homoptera). Part 1. Trans. ii. ent. joc. Lond. ^ : 47-60. 12. FtfiiWiAh, P. 1940. A contribution to the study of the genus Jfezara iunyot et Berville (hemiptera: Pentatomidae). Trans. Koy. Lnt. 00c. 90: 351-374. 13# GiilTLbK, L. 1938. Ueber den Bau des ±iLdri, iT. 1956. Polyteny as a factor in the chromosomal evolution of the Pentatomini (Hemiptera). Chromosoma 8: 135-151• HUuH^ -SChjiAULiK, b. and F. 1957* iMezara complex (Pentatomidae: Heteroptera) and its taxonomical and cytological status. J. horpn. 101: 1-23* iM-ib, a.l>. i960, a general Textbook of entomology. Lona. pp.^4-465. JAi'Jjjiii, bmijm 1959a. An analysis of the chromosomes in the four species of the family celostomatioae (heteroptera, cryptocerata). i. I960. Pre-reductional sex chromosomes in the family i'ingidae (Gymocerata-heteropteraj. £he Nucleus 3 (2) 209-214. KlKttALDY, G.w. 1906. Home remarks on the phylo^eny of the hemiptera-heteroptera. Can, i^ nt. 40: 357-3b4. KIRKaIjuY , U.'.rf. 1909* Gatalogue of hemiptera (heteroptera) I. Cimicidae Berlin Airi-392 pp. University of Ghana http://ugspace.ug.edu.gh KPOKDUOBiii KUMAR, R . KUWaH , R . K U im R , R . KUi-iAK , R . KUaiaR , K . Ljbibi'UN, i) , P.G.O. 1979. Karyotypes of some tropical reduviids and cytotaxonomy of Reduviidae. (Hemiptera: Heteroptera). h.bc. Ihesis Univ. of Uhana 146 pp. 1962. Morpho-taxonomical studies on tne genitalia and salivary glands of some Pentatomoidea. iunt. Tidskr Stockholm 83: 44-88. 1964. On the structure and function of the so-called ejaculatory reservoir in Pentatomoidea (Hemiptera: Heteroptera). Proc. Roy, froc. d^. 75: 51-65* 1965. Contributions to the morphology and x’Celationsnips Pentatomoidea (Hemiptera: neteroptera) Pt I. scutelleridae. 0. mit. froc. d^. j4, 41-55* 1968a. Aspects of the morphology and Relationships af tne superfamilies lygaeoidea, Piesmatoidea and Pyrrhocoroidea (Hemiptera: Heteroptera). .ant, hon. Mag. 10^ : 251-261. 1968b. morphology and Relationships of the Pentatomoidea (Heteroptera) IV. oncomerinae (Tessaratomidae). Aust. cJ. zool. 17: 553“696. 1971. Morphology and Relationships of the Pentatomoidea (Heteroptera) V. Urostylidae. i'he Arner. 1-iidland Naturalist. _1_; 63-73. 1952. Notes on the Ethiopian Pentatomoidea (Hemiptera) VIII. Scutellerinae leacn of Angola, with remarks on the genitalia and classification of the subfamily. Pupl. cult, cia Uiamant. Angola l b : 9-26. University of Ghana http://ugspace.ug.edu.gh 34. LfckTuN, Dm 1954a* Wingvenation and genitalia of lessaratoma Berthold, with remarks on Tessaratomini Btal (Hemiptera: Pentat omidae). Proc. K. n^t. Boc. Lond. 2£: 9-16. 35. Lck/i'uN, 'Dm 1954b. The male genitalia of behirus bicolor (L) (Heteroptera: Cydnidae) U. Boc. Brit* ant. 5_: 75*78. 36. LBbTOW, Dm 1955. A key to the genera of Oncomerinae Btal (Hemiptera: Pentatomidae) Proc. H. ent* Boc. Bond. 24: 62-68* 37. LiiioTOM, D.,1958* Chromosome number and the systematics of Pentatomorpha (Hemiptera; Proc. 10th int. Congress. Bnt. Montreal 2_: 9 H “9l8. 38. LtiioTuM, Dm, PMDjCiRGioc. Bengal J>: 155“l6l* 44. Maxima, G.K. 1951. A study of the chromosomes during meiosis in Forty-three species of Indian Heteroptera. Ibid. it: l-llb. 45. MAMma, G.K. 1958. Cytology and Interrelationships between various groups of Heteroptera. Proc. 10th Int. Congress lint. 1956 2: 919-934. 46. MaaT IN , B .A . , 1953* Temporary elimination of the autosomes from the meiotic spindle in a Halycinid Pentatomid. J. Morph. Philad. 92: 207-239* / 4?. MIYAMOTO, o. 1957* List of ovariole numbers in Japanese Heteroptera. bieboldia. Fulkuska 2.: 69-82. ^48. MIYaMOTo , ti. 1961. Comparative morphology of the alimentary organs of Heteroptera with the phylogenetic consideration. sieboldia 2(4), 197-259. 49* MONTGOMERY, T.H. 1901. Further studies on the chromosomes of the Hemiptera, Heteroptera. Proc. Acad. Nat, ^c. Philad. 53.: 261-270. 50. MONTGu j^lRY, T.H. 1905. The spermatogenesis of £>yrbula and Lycosa with general considerations upon reduction and the hetero chromosomes. Ibid. 57: 162-205* 51. MuNTGORuKi, T .H ., 1906. 1. Chromosomes in the spermatogenesis of the Hemiptera. Trans, i-uner. Jrhil. o^c. 21: 97-173. University of Ghana http://ugspace.ug.edu.gh MONTGOMERY, T.H., 1906. 2. The terminology of aberrant chromosomes and their behaviour in certain Hemiptera, Science 23* 36-38« QKBaLa , T. 1947* On the formation of bivalents in some Pentatomid bugs (Heteroptera) Hereditas. Lond. 33:110*118. OwUbU-MANU, L., 1971* Bath.ycoelia thalassina, another serious pest of cocoa in Ghana. C.M.B. Newsletter. 47; 12-14. PaRBHaD, R. 1937a. Chromosome number and sex Mechanism in Twenty species of the Indian Heteroptera. Current science 26: 123• PaR£HA.u, R. 1957b. Cytological studies in Heteroptera. III. A Comparative study of the chromosomes in the male germ-cells of eleven species of the subfamily Pentatominae. Res. Bull. Panjab. Univ. 122: 401-420. PiuNi>iiiRGKAbT, J.G. 1937* studies on the reproductive organs of Heteroptera with a consideration of their bearing on classification. 'Trans. R. ^nt. Boc. Lond. 109: 1-63. PRUTHi, H 1 9 2 3 . 'The morphology of the male genitalia in Rhynchota. Trans, -snt. soc. Lond. 77: 127-26?. RAO, s.R.V. 1934. Meiotic studies in three species of Heteroptera. bool, soc. India. Calcutta 0: 153-158. BCHaCHuw, B.jj. , 1932. Material Uber die chromosomen der Hemiptera. Anat. Anz. Jena 73: 456-479. University of Ghana http://ugspace.ug.edu.gh SCHAiiiFKR, C ♦SGHAEFbLR, ¥ oGHOUTlSDEN, *oCHOUTEDENf SGili^AjJiW 9 F SGHRajJMi, F SCHRADER, F SCHRAD.bR , F .W. 1964# The morphology and higher classification of the Coreoidea (Hemiptera-Heteroptera): Parts I and II. Ann, ent. o^c. Amer. 21} 670-684. ;.W. 1966. Some notes on Heteropteran trichobothria. The Michigan Entomologist 1 (3): 83-89. n. 1905. Faune entomologique de l*Afrique tropicale II. Arminae et Tessaratominae. Annls. Hus, r. Gongo. Belg. ser., 8 ve (2) 1. 133-277* H. 1910. Sjostedts Kilimandjaro-Meru Expedition 12 Hemiptera 6 Pentatomidae. 73-96. 1943a. Regular occurrence of heteroploidy in a group of Pentatomidae (Hemiptera). Biol. Bull. Lane. Pa 88: 63-70. 1943b. The cytology of regular heteroploidy in the genus Loxa (Pentatomidae-nemiptera). J. norph. Phild. 26: 157-177. 1946a. The elimination of chromosomes in the meiotic divisions of Brachystethus rubromaculathus .uallas. Biol. Bull. Lane. Pa §0: 19-31. 1946b. Autosomal elimination ana preferential segregation in the harlequin lobe of certain Discocephalini (Hemiptera). Biol. Bull. Lane. Pa. 265-290. 1947. The role of the Kinetochore in the chromosomal evolution of the Heteroptera and Homoptera: University of Ghana http://ugspace.ug.edu.gh Evolution 1_: 154-142. 70. F. I960. Evolutionary aspects of oberrant meiosis in some Pentatominae. (Heteroptera) Evolution Lane* 14: 448-508. 71. SGHRauiuK, F. and HUuHBS-SCHJSAJiiR, b. 1956* Polyploidy and Frag­ mentation in the chromosomal evolution of various species of Thyanta (Hemiptera). Ghromosoma 7_* 469-496. 72. bCUUUliLtt, G.C.L. 1959. 'The female genitalia of the Heteroptera Morphology and bearing on Classification. Trans. K. Lnt. froc. Lond. Ill: 405-46?• 73* SHaRHa , G.P. and PaRSHa.u,R. 1955* '^he morphology of chromosomes in Laccotrephes maculatus (Fab.) (Heteroptera) Hes. Bull. Pan .jab. Univ. 72: 62-72. 74. SINGH, iuft and SINGH, 6.0. 1966. Meiotic studies in the bug. Chrysocoris stollii wolff. Natur wissenschaften 53: 91 75* SIWNaDUKAI, P., 1979* a revision of the Dinidoridae of the world (Heteroptera: Pentatomoidea) Keys to, ana descriptions of subfamilies, tribes, genera and species, with designation of types. Ph.D. thesis. University of Ghana, Legon. y r 76# SOUTHWOOJJ, T.fi.ii. 1956. The structure of the eggs of the terrestrial Heteroptera and its relationships to the classification of the group. Trans. Roy. unt. boc. Lond. 108: 163-221. University of Ghana http://ugspace.ug.edu.gh 77. SlIS, P. 196^ . Thausmastellidae - a new family of pentatomoid Heteroptera* Acta boc* ent. Csl. 61: 238-253• 78. TGSHIOKA, S. 1936. On the chromosomes of some Hemipterous Insects* Cytolo&ia* 7* 403-^ 06. 79. IESHIMA, N. 1963. Chromosome study of Thyanta Pallidovirens (Stal) in relation to Taxonomy (Hemiptera: Pentatomidae). Pan* Pacific Ent. 39 (3): 1^ 3• 80• WA&REN, E.M., GASSNEH, G. and Bj3ELa1nI1j, u.P., i960. Somatic chromosomes of praying mantids as shown by the squash techniques. Tech* Reports on Biology and Medicine 18: 67^ 678* 81• WHITE, 1956. Cytogenetics and systematics Entomology. Ann. Rev. Ent. 2_: 71*90* 82* WILSON, iii.B*, 1903a. Studies on chromosomes I* 'The behaviour of Idiochromosomes in Hemiptera. J. Exp. bool. 2: 371-^5 • 83. WILSON, E.B. 1905b. Studies on chromosomes II* The paired micro chromosomes, idiochromosomes and heterophic chromosomes in Hemiptera. J. Exp. aool. Zi 507-5^5* WILSON, E.B. 1906a. 'The chromosomes in relation to the deter­ mination of sex in insects. Proc. Soc. Exp* Biol* Med. 3: 19-23. 85* WILSON, E.B. 1906b. Studies on chromosomes III. The sexual difierences of the chromosome groups in Hemiptera with some considerations on determination and inheritance of sex. J. Exp. Eool. Med. 3; 1-^ 0* University of Ghana http://ugspace.ug.edu.gh 86* WILSON, i!J.B. 1907. The case of Anas a tristis. Science 25* 191-193- 87o WILSON, K.B. 1911. Studies on chromosomes. A review of the chromosomes of Nezara, with some more general considerations* J* Morph. Phild. 22: 71-^“lO# 88* WOOJJwiUCD, T.L. 1950. Ovarioles and testis follicle numbers in the heteroptera. Lnt. lion, hag, -^ ond. 86: 82-84. 89# VaN L.p* I917. Heport upon a collection of Hemiptera made by waiter M. Clifford in 19^ 6 and 1917- Proc. Calif. Acad. Sci. 11: 249-318. 90* YQsId a, T.H. 1956. Studies on the chromosomes of coleopteran and Hemipteran insects, with special regard to the (Juantitative relationship between autosomes and sex chromosomes. Proc. 10th int. Congr. ^nt. Montreal 2: 979-989• * original not seen University of Ghana http://ugspace.ug.edu.gh APPENDIX 'jtABUi 1 LIST OF SP&CUjo OF PriNTATOMOIJJEA USiCii A5 MaTjJHIAL SPLCjSIS LOCALITY ------- .— - ------------- DaTL OF COLLECTION Acrosternum heegeri i‘ieb. Legon (UV).(S),S«/eija(S) May, June, July, August, 1979 Aeliomorpha sp.griseoflavs Aburi(S), Legon(S), May-August, 1979 WeijaUO Aeptus singular is (Dali.) Legon(S) September, 1979 Aethemenes chloris (westw.) Legon (UV). March, 1979 Aspavia acuminata Mont. Legon(S), Tafo(S) July, uec ember 1979 Aspavia armigera (jfabr.) Legon(S), humasi(b), May, July-Beptember, Kade(b), weija(o) December, 1979 Tafo(o) Aspavia hastator (ii'abr.) Kade(S),Aburi(S), ) Kibi(S) ,Mampong(S) i February, May-Oct. Tafo(s), suhum(s) 1979 Atelocera s errata (JTabr.) Tafo(S) December, 1979 Bathycoelia rodhaini Hade(UV), Tafo(HP) February,June 1979 Schout. Bathycoelia thalassina Le gon ( U V ) , Taf o (HP ), £‘ebruary-dune, 1979 (H.-S) January, I960 Benia sp.A Aburi(B) October, 1979 Carbula capito St§l Suhum(b), Aburi(S) May, June, July, September, 1979 Carbula carbula (Dist.) Aburi(S), Mampong( B ) January, 1980 Carbula marginella Stal Aburi(b), tade(s), July-December, 1979 Degon(^ ) Carbula melacantha btSl Amonokrom(B), Kibi(B) July-Detober, 1979 Taf0(0) Carbula sp. nr. sjostedti Aburi^o), Kumasi(s) October, December 197< Schout• Tafo(B) Caura pugillater (i'abr.) Kade(B), Tafo(s) October, December 197$' • University of Ghana http://ugspace.ug.edu.gh APPENDIX TaBLE 1 (cont ,d) LIST OF SP^CILoOF PENTATOMOIDEA USED Ab MATERIAL SPECEIS Diploxys bipunctata (*myot et berville) Amaxosana punctata i-tist. Durmia haedula bt8l Durmia lutulenta bt§l Durmia sp. Dymantis grisea Jen-Haar Eysarcoris inconspicuus (H.-b) Halyomorpha annalicernis (sign.) Halyomorpha retlexa (bign Halyomorpha picus Dist. Lenda punctata (Palisot-Beauvois) hacrina juvenca (burm.) mcrorhaphis acuta (Dali. Nezara viridula Linnr. Nezara viridula var smaragdula (Fabr.) Nezara viridula var torquata Puton. Antestia sp. iintestia sp.b.l nntestia sp. V immunda Linnr. antestiopyjs sp. o.l 1 LOCALITY Legon(UV),(b) Legon(b)(UV) Legon(b) (UV), i'afo(b) Kade(b), Legon(b), Tafo(b) Legon V^)(b) Legon (UV), ^umasi(s ) Legon (b) Tafo(s), j\ade (s) ) Bunso(b), Tafo(b) buhum(b) iiburi(b), Kade(b) hibi(s), i‘afo(s) buhum(s) Kade(b), Tafo(b), Kibi(b) ) Legon(b), Aade(b) Dodowa(b), Legon(s) ftade(b) Dodowa^s), iiegon(b) Dodowa(s), Legon(b), Legon (UV) Legon (UV) Legon (UV) legon (UV) DATE OF COLLECTION June, august, 1979 June, 1979 April, hay, 1979 July, August, 1979 June, 1979 august, September, 197'^ August, beptember,1979 November, 1979 January, 19o0 December, 1979 May, 1979 February, June-Octber, January, 1980 February-beptember, 1979 July-september, 1979 July-i>ec ember, 1979 January, I98O July, December, 1979 January, I98O February, July, Dec­ ember, 1979 iiay-July, I979 my-July, I979 May, June, 1979 May, 1979 University of Ghana http://ugspace.ug.edu.gh APPjSHCDIX TABLE 1 ( c o n t 1 d ) L IST OF SPujCILo OF PLNTATuHOIjJFA USEjj AS MaT^KIAL SPLCFIS LOCALITY P ie z o d o r u s h y b n e r i (G m e lin ) B c o t in a p h a r a f i b u l a t a & c h o u t. obepontia misella Stal Tyoma v e r r u c o s a M o n t. Veterna sanquineirostris Stal Afrius purpureus ( to e s tw . Fai'iILY SCUTTfoLi.i^ kIjjaL Callidea duodecimpunctat< l StRl Hotea subfasciata (westwood) Sphaeracoris testudogris i i t S l Steganocerus multipuncta (de Geer) FAi-'ilLY BRaCHYPLATIual Brachyplatys incertus (walker) Bracnyplatys testudoni« gra(.be Geer) Coptosoma nubila (Germ.) Legon(S), Lodowa(S) Kade(SJ Aburi(s), iiade(b), Suhum(S), Tafo(b) Aburi(S), Tafo(o) Kade(s), Tafo(s) Legon(S), Tafo(S) Asutuare (BP), -Legon Aburi (s) , ilibi (S) , Suhum(o) Legon(s) a Kade(i>), Tafo(b) Tafo(S), Kade^ S) K a d e ( s j , T a f o ( b ) K u m a s i(b ) L e g o n ( b ) , T a f o ( s ) DATJsi OF COLLECTION July-September, 1979 November, 1979 February-i>ec ember, 197 ? September, October, 1979 February, 1979 January, 19&0 September, December 1979 (4p)February-uecember,197< January-^pril, 19&0 September, November 1979 September, 1979 November, December 1979 November, December 1979i January, 1980 November, jJecember, January, 1980 Au^ust-oeptember,1979 November, 1979 University of Ghana http://ugspace.ug.edu.gh A PP J^uU . TaBL& 1 ( c o n t ’ d ) L IST OF SP^C IL o OF PLNTATOMOILjEA USlSi) AS MATERIAL SPLCJiilS LOCALITY 1-- — ------- — ----- DATiiJ OF COLLECTION Coptosoma stali Mont* Legon(b), Aburi(b), August-November, 1979 Tafo(b), ftade(b) January, 1980 family DiwiDoniDAii) Coridius cuprifer (westw.) Tafo(b), Kade(&), Abur i-(b) September, 1979* January, 1980 Fa^IlY TlttiSSAKATuMIDAjai Piezosternum calidum Tafo(b), Kade(o), (otal) Aburi(b) oeptember-Lecember, 1979 January, March, I9S0 FAMILY CYMIDaL Macroscytus sr». Legon (HP) (LTV) Feoruary-July, 1979 *=>eptember-itovember 1979 l-jay, 1980 (b) = Sweeping (HP) s Hand picking (Uv) = Ultra violet trsp University of Ghana http://ugspace.ug.edu.gh APPENDIX. Ta E.lE 2 SUMMARY OF HB3ULTB n a m e m a l e 2N NUHHtUR KAKYO'fYPE FaaviI1jY Pii^lATGMlDAE Acrosternum heegeri Fieb. Ik 12a +A+Y Aelioraorpha sp.? griseoflava btal Ik 12A+a +Y Aeptus singularis (Dali.) Ik 12 a+a +Y Aethemenes chloris (toiestw.) Ik 12a +a +Y Afrius purpureus (westw.) Ik 12a +a +Y Aspavia hastator (Fabr.) Ik 12A+A+Y Aspavia armigera (Fabr.) Ik 12A+a +Y Aspavia acuminata aont. Ik 12A+X+Y Atelocera serrata (Faor.) Ik 12a +X+Y Bathycoelia thalassina (.-o) Ik 12A+A+Y Bathycoelia rodhaini ochout Ik 12a +a +Y Benia sp.A Ik 12A+A+Y Carbula capita otal Ik 12a +a +Y Carbula carbula (Dist.) Ik 12a +a +Y Carbula marginella otSl Ik 12A+A+Y Carbula melacantha i^tSl Ik 12a +a +Y Carbula sp. nr. sjostedti ochout. Ik 12«.-*-a +Y Caura pugilator (Fabr.) Ik 12A+A+Y Diploxys bipunctata (Amyot et oerville) Ik 12A+A+Y Amaxosana punctata Dist. Ik 12a +a +Y Durmia haedula btSl Ik 12a +a +Y Durmia lutulenta stSl Ik 12a +X+Y Durmia sp. Ik 12a +a +Y uymantis grisea oen-Haar Ik 12a +X+Y Eysarcoris inconspicuus Ik 12A+a +Y Farnya vessicolor (Fabr.) Ik 12a +a +Y Halyomorpha annulicornis (M 'i Ik 12a +a +Y University of Ghana http://ugspace.ug.edu.gh APPEiMlXU I'AJiLE 2 SUMMARY UF KEBULTb(cont'd) NAME MALE i 2N NUMBER KAKYOTYPE Lalyomorpha picus (uist.) 14 12A+A+Y tialyomorpha reflexa (^ i^ n.) 14 12A+A+X Lerida punctata (Palisot Beauvois) 14 12a+A+Y A^ iacrina .iuvenca (Burm.) 14 12A+A+Y Macroshaphis acuta (i/all.) 14 12A+A+Y Nezara viridula (L.) 14 12A+X+Y Nezara viridula var smara^dula (Fabr.) 14 12a+A+Y Nezara viridula var torquata (Puton) 14 12A+a+Y Antestia so* 14 12a+a+Y nntestiopsis sv. b* I 14 12ii+X+Y Antestia sp* o. I 14 12h+a+Y Antestia su. ? immunda liinnr* 14 12a+a+Y Piezordorus hybneri (Gmelin) 14 12A+A+Y Bcotinophora fibulata (bchouteden) 14 12A+A+Y ^epontia misella btSl 14 12a+a+Y 'lyoma verrucosa (Mont.) 14 12a+a+Y Veterna sanquineirostris btSl 14 12ii+A+Y FAMILY bCUTELLEKlJJAE Gallidea duodecimmmctata ) 12 lOii+A+Y notea subfasciata (westw.) 12 lOii+A+Y Bphaeracoris testudoep?isea StQl 12 10a +^ .+Y bteganocerus multipunctata (Hp ( W yO 12 10ii+A+Y University of Ghana http://ugspace.ug.edu.gh ICABLiii 2 SUMMARY UF RESULTS (Cont' d) NAME MaLE 2N NUMBER KARYOTYPE FAMILY BRACHYPLATIDaE (PLAIASPIDaE) Brachyplatys incertus (walker) 12 IOa+A+Y Brachyplatys testudo nigra St§l 12 10a+a+Y Coptosoma nubila Uerm 12 10a+a+Y Coptosoma stali (Mont*) 12 10j*+a+Y FAMILY DINlDGRIDAE Coridius cuprifer (Westw*) 14 12A+A+Y FAMILY TEdSAKATGhIDAE Piezosternum calidum btSl 14 12a+A+Y FAiviILY CYNlDAE idacroscytus sp. 12 10a+a+Y University of Ghana http://ugspace.ug.edu.gh APPm>IX TABLE 3 CYTOUJGICaL WORK ON PjbNTA'ruMUlDJWt. FROM LI'JbRKl'UKk. Family Subfamily No of J Species with known 2n 2n number ACAN ,£HGSQi'/iATIDAE 6 IOa+X+Y = 12 'IciSaAl^ TOMIDAE 3 10A+X+Y = 12 Tessaratominae Natalicolinae j Oncomerinae 2 1 10A+X+Y = 12 IOa+X+Y = 12 CYDNLDA^ tthirinae 7 6 = 10a+a+Y = 12 1 = 21A+A+Y = 51 SCU-mJii-aiDAE bcutellerinae Pachycorinae Eurygastrinae 15 10a+a+Y = 12 7 1 7 IOa+a+Y = 12 IOa+a+Y - 12 10a+a+Y = 12 E^CHYPLaTIDAE Coptosominae 8 10a+a+Y = 12 DINIDGKIDAE Dinidorinae 6 3 = 12A+X+Y = 14 1 = 18A+A+Y = 21 2 = 18a+a+Y = 20 EUmiWOIIDAE 1 12A+X+Y = 14 UHObTYLIDa E — ■ ■ . A________ 2 14a+a+Y = 16 University of Ghana http://ugspace.ug.edu.gh APPiiNDlX TABLE 3 (cont'd) 105 Family Subfamily --------rrr No of Species with known 2n --------------------- 2n number t P^'Ta TGMII)A£ i I 170 1 = 4A+X+Y = 6 1 = 8A+A+Y = 10 4 =10A+X+Y = 12 L42 =12a+a+Y = 14 1 =12A+2X+Y= 15 14 =l4ii+X+Y = 16 6 =24a+X+Y = 26 1 =24a+2a+Y=27 POIXJPINA& 1 12A+X+Y = 14 PHYLLuCEPHAEliiaL - - a MYOTEINa E 14 10 = IZa+A+Y = 14 3 = 14a+a+Y = 16 1 = 10a+a+Y = 12 PEN Ta TUMIN a E 155 1 = 4a+a+Y = 6 1 = 8a+a+Y = 10 3 =10A+a+Y = 12 131 =12a+a+Y = 14 1 =12A+a+Y = 15 11 =14a+a+Y = 26 6 = 2 W + I = 26 1 =24a+a+Y = 2? § University of Ghana http://ugspace.ug.edu.gh APPifliL»IX lEABLni k 106 0IS; TloN OF 2 li^i THiij FAMlLXLo OF PLNlAlOMOI-UbA STUiJXj&J CYTOLUilCiiLL Y TO DtiHh Family Subfamily No of Species with known 2n 2xi number ACANTHOSOMATIDiift 6 6 = IOA+a+Y = 12 'Bl^ bbAitA'l'OHIBAE k 3 = 10a+a+Y = 12 1 = 12A+A+Y = Ik Tessaratominae 2 2 = 10A+X+Y = 12 Watalicolinae - - Oncomerinae 2 1 = 10a+a+Y = 12 1 = 12A+X+Y = 14 CYDNIDAE Sehirinae 8 7 = 10A+X+Y = 12 1 = 28a+2a+Y= 31 SCUT WL1 iftRIDAE 18 18 = 10A+X+Y = 12 Scute Her inae 10 10A*X+Y = 12 Pachycorinae l 10A+a+Y = 12 Eurygastrinae 7 IOa+X+Y = 12 BKaCMYPLATIDAS COPTGbOMIWAitl 10 10a+a+Y = 12 DliMTDOHIUAL D11'U1)0kIInIa£ 7 l+ = 12A+A+Y = Ik 1 = 18a+2a+Y= 21 2 = 18a+a+Y = 20 EUMatfOTlDAJai 1 12A+A+Y = Ik UHOSTYLIDAJB 1 14a+a+Y = 16 PjiNTATOMlDAiii 211 1 = 4a+a+Y = 6 1 = 6a+a+Y = 10 k =10A+a+Y = 12 183=10A+a+Y = Ik 1 = 12a=2X+Y = 15 University of Ghana http://ugspace.ug.edu.gh APPi^D lX T a b le 4 ( c o n t 1) 107 Family Subfamily No of Species with known 2n 2n number PisJNTATOMIBAE 14 = 14a+a+Y = 16 6 = 24a+a+Y = 26 1 = 24a+2X+Y= 2? PODOPINAE 1 1 = 12a+a+Y = 14 PHYLLOCi^ HjiLlNAiii 1 1 = 12a+a+Y = 14 AMYOTLINAiC 15 1 = 10a+a+Y = 12 11 = 12a+a+Y = 14 3 = 14a+.v+Y = 16 P&N TATuMN Aiii 194 1 = 4a+a+Y = 6 1 = 8a+a+Y = 10 3 = 10a+a+Y= 12 170 = 12A+a+Y= 14 1 = 12a+2^+Y=15 U = 14a+a+Y =16 6 = 24a+a+Y = 26 1 = 24a+a+Y b 27 University of Ghana http://ugspace.ug.edu.gh APPflMDlA TABLE 5 nT S 'P B T H U 'riuM UF 2 M ftlUI-JBMto IN 'X'Jdb b l A grtiH l-U LLa O F Pi^MTAX'OhOlULA WOKALU OIM THIS 103 Family Subfamily Frequency 2n Number 'fessarat omidae Oncomerinae 1 12A+A+Y = 14 Scutelleridae Scutellerinae k IOa+a+Y = 12 Dinidoriuae Dinidorinae 1 12A+X+I = OA Brachyplat i dae Coptosominae 4 lOii+A+Y = 12 Pentatomidae Amyoteinae Phylloc ephalinae Pent at ominae 1 1 42 12a+a+Y = 14 12a+a+Y = 14 12A+a+Y = 14 CYDNIDaE Sehirinae 1 10a+a+Y = 12 University of Ghana http://ugspace.ug.edu.gh APHjNDIX TABLE 6 C y to lo g ic a l work on Acanthosom atidae from L i t e r a tu r e f am il y Acanthosomatidale Acanthosoma denticoada Yakoulev Acanthosoma haemorrhoidale (Lin) Acanthosoma labiduroides Yakouleve SPECIES Elasmastethus humeralis Yakoulev Elasmostethus interstinctui IOA+a+Y (Linn*) Elasmucha securvum Dallas 2n lOa+iC+Y IOa+a+Y 10a+a+Y 10a+a+Y 10A+X+Y reference and LOCALITY Of MATERIAL Miyamoto 19579 Japan Woodward 19509 England Yosida 1956, Japan Yosida 1956, Japan halkka 1956, Finland Parshad 1957b,India * Locality of material under­ lined. University of Ghana http://ugspace.ug.edu.gh aPPLNDIa TABL& 7 Cytological work on Tessaratomidae, Cydnidae from literature family SPECIES / 2n REFERENCE AND LOCALITY OF MATERIAL* Tessarat omi dae I'essarat ominae Eusthenes saevus btSl 10a+A+Y Parshad 1957b, India i Tessaratoma qauanica IOa +a +I Parshad 1957b, Thunberg Subfamily Qncomerinae Piezosternum subulatum bt8l 10a+a+Y -Schrader 19^71 Brazil Family Cynidae Sehirinae Sehirus bicolor (Linn.) IOa +a +Y Leston 195^ b, U.il. \ Parshad 1957b, j India if Stibaropus molgimus btSl 28a+a+Y ii Aethus maurus Dallas 10A+a+Y Jande 1959b, India it Aethus varians (Fabr.) 10a+a+Y Jande 1959b, India Jande 1959b, India Kumar (Unpublished) Ghana ii Aethus ni^ritus (Fabr.) 10a+X+Y ii Macroscytus brunneus Stal 10a+a+Y it hacroscytus subaenus (Fabr.) IOA+a +Y Parshad 1957b, India * Locality of material under­ line. University of Ghana http://ugspace.ug.edu.gh APPENDIX Ta IILL 8 C y to lo g ic a l work on S c u te l le r id a e from l i t e r a t u r e FAMILY SPECIES 2n reference and LOCALITY OF material * f SCUmLiSKIDAE Subfam* ! &cut ellerinae Cantao ocellatus Thungberg 10a+a+Y Banerjee 195$, India it Chrysocoris eques Distant 10a+a+Y Jande 1959b,India if Chrysocoris stollii Wolft 10a+a+Y Singh and Singh Japan ii Scutellera nobilis distant IOa+^+Y Jande 1959b, India i Scutellera perplexa (westw. ) 10A+a+Y Hanna 1958b, India j ii Solenostethnim rubropunc- tatum Distant 10A+a+Y Janae 1959b, India it Subfamily Sphaerocoris testudogrisea (de Geer) 10a+a+Y Kumar(Unpublished) | f Ghana Pachycorinae Subfamily Hotea curculioneides Vollenhoven IOA+a+Y Manna, 195$* India j !! i Eurygastrinae Lurygaster austriacus Schrank IOA+a+Y Montgomery, 1901, I 1906, U*0•A• ii Lurypaster alternatus Uhler 10a+a+Y Schachow 1932, , Germany ! ii Lury^aster hotlentota 10A+a+Y Schachow 1932, Jakoulev Australia t LuryRaster maura maunders 12A+A+Y Geitler 193&, Australia,Germany ii Lurygaster maura baunders 10a+a+Y schachow 1932, Australia ii p.dontat arsus caud^tus Horvath 10a+a+Y ochachow 1932, Germany \ -----— | * Locality of ma­ terial underlinec• University of Ghana http://ugspace.ug.edu.gh aPPj^ mpIa TaBLK 8 (cont'd) Cytological work on Scutelleridae, Brachyplatidae from literature fam ily SPECIES 2n REFERENCE AND LOCALITY or; MATERIAL Scutelleridae gastrinae Gdontotarsus purpureo- 10a +X+Y Schachow 1932, lineatus norvath Germany i f Odontotarsus robustus 10a +a +Y Schachow 1932, Jakoulev GermanyBKACHYPiiiiTljj^ Brachyplatys pauper IOa +a +Y Kao 195^ , India (Vollenhoven) i t Brachyplatys subaeneus 10a +X+Y Manna 1958, India, (i/i/estw.) Banerjer 1958, India i t Brachyplatys testudoni^ra 10a +a +Y Kumar (Unpublished) total Germany i i Coptosoma biguttula IOa +a +Y Yosida 1956, Japan (Motschlsky) i t Coptosoma cribarium (tfabr.) 10a+a+Y Banerjee 1958, India i i Coptosoma punctissimum(Mont ►)IOa+^+Y Yosida 195°, Japan i i Coptosoma stali (hont.) 10a+a+Y Kumar (Unpublished) Ghana t i Coptosoma varie^ata (H.-6) 10a +a +Y Jande 1959b,India * Locality ofm a t e r i a l u n d e r ­l i n e d . i University of Ghana http://ugspace.ug.edu.gh APPENDIX ‘lABLti 9 Cytological work on Ldnidoridae from Literature family SPECIES 2n i REFERENCE AND LOCALITY OF MATERIAL * Dini dor i dae Goridius .janus 12A+.A.+Y Manna 1958, India (Aspongopus) Fab. i i Goridius orientalis 12A-J-A+Y Manna 1951, India (Aspongopus) Kirkaldy i i Coridius sp 12A+X+Y Jande 1959b, India i i Dinidor ruficocinctus (btal ) 18a+a+Y Schrader F. 19^ 7* IndicL i i Megymenum brevicorne (Dalla 3)18a+X+Y Banerjee 1958, j India 1V i i Megymenum gracilicorne l8ii+A+Y Schrader i960, j Costa Hica j * Locality of \ material under- f lined* 1• \ i I j \ \* ) i 1 University of Ghana http://ugspace.ug.edu.gh APPENDIX TABUS 10 C y t o lo g ic a l work on Pen ta tom idae from l i t e r a t u r e family SPECIES 2n REFERENCE AND LOCALITY GEm MATERIAL' Pentatomidae Pent at ominae Ablaptus amazonus StSl 12A+X+Y Schrader I960, Costa Rica it Acrosternum hilasis bay 12A+X+Y Montgomery, 1901 y «s•a * ti Acrosternum marginatum(P&B) 12A+X+Y Hughes-Schrader .and Schrader, 1950 Panama* ................. V ti Acrosternum sp 12A+A+Y Hughes-Schrader and Schrader, 1957 Trinidad it Acrosternum pennsylvanicum (P. and B.) 12A+a+Y Hughes-Schrader and Schrader, 1957 %USA ti Acrosternum scutellatum Distant 12A+A+Y Hughes-Schrader and Schrader, 1957 Costa Rica i i Adria parvula Ber^roth 12a+^+Y . Jande 1959» India it Aelia acuminata (Linnaeus) 12A+A+Y Schachow 1952, Germany i i Aelia fieberi Scott 12A+A+Y Miyamoto 1957•Japan i i Aelia rostrata Boheman 12a+a+Y Yosida 1956, Japan i i Aelionorpha griseoflava stSi 12A+A+Y Kumar, unpublished Ghana i t Aenoria lewisi (^ cott) 12A+a+Y Jande 1959b, India i i Agach.itus dromedanius btSl !2ii+A*hY ^chraaer I960, Costa Rica * locality of > material under­ lined. University of Ghana http://ugspace.ug.edu.gh APPLNdIX TABLE 10 (GOi'JT'u) f am il y Pentatomidae Bub Pentatomina' SPECIES Agonoscelis nobilus (Fab.) Alitocoris parvus (distant) Alitocoris schracteri(Sailer) 12a +a +Y Ancyrosoma alliolineata Antestiopsis cruciata (Antestia) Architas pudeus (dist.) ArVeluis albopunctata (de Geer) Aspavia ingens distant Bagrada picta (Fabr) Banaso bidens schraderi Bailer Banasa celva (Bay) Banasa centralis Bailer Banasa dimidiata (Bay) Banasa enchlora (btal) 2n 12A+X+Y 12A+X+Y 12a+a+Y 12A+A+Y 12A+A+Y 12A+A+Y 12A+A+Y 12A+A+Y 24a+a+Y 24a+X+Y 24a+A+Y 14a +a +Y I^A+a+Y REFERENCE AND LOCALITY OF MATERIAL’" Manna 1951? India Schrader I960, Panama Schrader I960, Costa Rica Schachow 1932, Germany Manna 1951* India Schrader i960, Panama Hughes-Schrader and Schrader 1956, Cost* tdca Leston unpbl., Ghana tiao 1955» Mitt.India Schrader and Hughes1 ■ bchrader 1956, Cost, i Rica, Panama Wilson, 1905, 1907, Schrader and Hughes-* Schrader 1956,U»S»a Schrader and Hughes- Schrader, 1956 Costa Rica Wilson 1907; Schra­ der and Hughes- bchrader 1956,USA Schrader and Hughes- ochrader 1956,U.a.a University of Ghana http://ugspace.ug.edu.gh APgiiMDIX TABLE 10 Ccont'd) FAMILY SPECIES 2n REFERENCE AND LOCALITY OF MATERIAL* Pentatominae Banasa lenticularis Uhler 14a+a+Y Schrader and Hughes- Schrader 1956,USA i i Banasa minor Sailer 24a+a+Y Schrader and Hughes- Schrader 1956,USA i i Banasa panamensis Sailer 12A+A+Y Hughes-Schrader and bchrader, 1956 ,1958 Panama i i Banasa rufifrons sailer 24a+X+Y Schrader and Hughes1- Schrader 1956 Panama t i Banasa zeteki sailer 24a+a+Y Schrader and Huahes • Schrader 1956, Panama n Brachystethus rubromacu- latus Dallas 12A+A+Y i bchrader I960, J Costa iiica i i Carbula biguttata Distant 12A+X+YX Yosida 1956, Japan i i i Carbula aspavia Distant 12A+A+Y Parshad, 1957b » ' Jndia j n Carbula socia Distant 12A+A+Y Manna 195o, India * i t Carbula sp. 12A+.W+Y Manna 1956, India ! t i Carpocoris melanocerus Mulsant et Key 12A+A+Y Ueitler 1958, Australia i i Carpocoris pudicus (Poda) 12A+X+Y bchachow 1952, Germany i i Captus pallipes (Neodius) 14a+a+Y Parshad 1957a,b, India i i Ghlorochroa juniperina (Pentatoma) 12*+a+Y Wilson 1905a, UsA i i Codophila varia 12a+a+Y schachow 1932, Germany i i Coenus delius (Sav) : 12a+.a.+Y 0 Montgomery 1901, University of Ghana http://ugspace.ug.edu.gh APPENDIX T aB IiE 10 ( c o n t 'd ) family SPECIES REFERENCE and LOCALITY OF MATERIAL* Pentat omidae Subfam. Penta Cosmopepla lintneriana (Kirkaldy) Dalpada confusa Distant Dalpada concinna (westwoodj Dalpada versicolor Distant Dinocoris baccarum (Pentatoma) Dolycoris indicus Stal Odessa caldaria Distant Odessa celsa Distant Edessa costae Bergroth Edessa fuscidorsata (Fabr.) Odessa irlorata Dallas 14A+X+Y 12A+a+Y 12A+a +Y 12A+A+Y 12a +a +Y 12a +a +Y 12a+a+Y 12A+A+Y 12A+A+Y 12A+A+Y 12A+A+Y I f Edessa laticornis btal 12A+A+Y I I Edessa pictiventris Stal 12a +x +Y I I Edessa rufomarginata ktal 12A+A+Y I I _Edessa vinula Stal 12A+A+Y I f Edessa so I 12A+..+Y I I iudessa sx> II 12A+A+Y I I Ddessa sp III 1 2 A + A + Y Montgomery 1901, 1906, U.S.A. Parshad 1957a, Indi|a Parshad 1957a * 11 Kao 1955» India Schachow 1932, Germany Manna 1951» India 'Schrader 1945a, Panama Schrader 1945a, Panama Schrader 1945a, Panama Schrader 1945a, Panama Schrader 1945a, Panama Schrader 1945a, Pgjfcga Schrader 1945a, ,f Schrader 1945a, Panama Schrader 1945a, Panama Schrader 19^ 5a, Panama Schrader 1945a, Panama ochrader i960, Panama University of Ghana http://ugspace.ug.edu.gh APPKMdIX TABLE 10 ( c o n t 'd ) FAMILY SPECIES 2n REFERENCE and LOCALITY OF MATERIAL* Pentatominae Lurydema dominulus Heuter &urydema festiva Hovarth ■burydema fieberi if'ieber Lurydema liturifera Distant n,urydema oleracea Hovarth Lurydema ornata Hovarth Lurydema pulchra distant iiurydema rugosa Motschulsky Lurydema ventralis Kolenati Kuschistus crassus Dallas Jiiuschistus fissilis Uhler fehischistus ictericus btal riuschistus obscurus Dallas j£urchistus serves Stal Luschistus tristigmus st§l ■kuscnistus variolarius stal Luschitus sp ^ysarcoris aenus (stollia) 12A+X+Y 12A+A+Y 12A+A+Y 12A+A+Y 12a +a +Y 12A+a +Y 12a +A+Y 12a +a +Y 12a +A+Y 10a +a +Y 12a +a +Y 12A+A+Y 12a +a +Y 12A+A+Y 12a +a +Y 12A+a+Y 12a +a +Y 14A+A+Y Geitler 1958f Australia Schachow 1952, Germany Parshad 1957a-,India Parshad 1957 * 11 Schachow 1932, Germany schachow 1932, Germany Parshad 1957a,India Parshad 1957a * 11 Geitler* 1938, Australia Mughes-schrader and schrader 1956, Usa Wilson 1905,1906,USA Wilson 1906, U.s»A Hu5hes-ochrader and Schrader 1956, USA Wilson 1906, U»s «a Montgomery 1906, Wilson 1906, U.s„A Montgomery 1906, Wilson 1906 U»s»A Montgomery 1906, U »s♦A,—■ schachow 1932, Germany University of Ghana http://ugspace.ug.edu.gh APPENDIX ‘I'Abiab 10 (c o n t * d ) family SPECIES 2n REl’ERENCE AND LOCALITY OF MATERIAL*- P entat ominae Eysarcoris capitatus Distan ; 12A+X+Y Manna 1951, India i i Eysarcoris guttiger Thunber 5 12a+a+Y Ma nna 1951? 11 i i Eysarcoris inconspicuus(H.s ) 12A+A+Y Parshad 1957*»M i i eysarcoris fabricii Hahn 14a+X+Y bchachow 1932, Germany i i eysarcoris montivaRns uistait 12a+a+Y Parshad 1957a, jjxdia i i eysarcoris parvus Uhler 12a+a+Y i'liyamoto 1957a, India i t iiiysarcoris ventralis (^ estwood) 12A+A+Y Miyamoto 1957*India t i Eysarcoris sp 12A+A+Y Miyamoto 1957, 11 i i Graphosoma italicuus (Muller) 12a+a+Y Geitler 1938, Australia i i Graphosoma rubrolineata (Westwood) 12A+A+Y Yosida 1956, Japan i i Graphosoma semipunctata (Fabr.) 12a+a+Y Schachow 1932, Germany i i Halyomorpha picus (i<'abr.) 12a+a+Y Parshad I957a,b, India n Halys dentatus Distant 12A+A+Y bharma and Parshad 1955, India i i Halys sulcatus 'fhunffbere 12a+a+Y Manna 195&« India i i Holcostethus limbolariusStSl 12a+a+Y Hughes-schrader and bchrader 1956, IfoA i t Lerida punctata (Palisot Beauvois) 12A+X+Y Leston Unpbl., Gnanc, i i Loxa flavjcolliR ( U r n e y '1 12a+a+Y Schrader 1545b, ^osta ideai i L o x a . florida (van 14a+^+Y ochrader 1945b rifc>A University of Ghana http://ugspace.ug.edu.gh APPi^U lX TABLfl 10 ( c o n t 1 d ) family SPECIES 2n REFERENCE AND LOCALITY OF MATERIAL * Pentatominae Macropygium reticulare 12A+A+Y ^chrader I960, Costa Kica it Mayrinia variegata (Distant ) 12a +^+Y Schrader 19 5^a, I960, Josta kica ti Menida ben&alensis (westw.J 12a +a +Y Banerjee 1958, India it Menida violacea (Notschulsky.) 12a +a +Y Yosida 1956,India it Melanoderma apicifera Distant 12A+/V+Y ochrader I960, Costa Kica it Mecistorhinus tripterus (j.-’ab. ) 12A+A+Y Schrader I960, Costa Kica j i i Mecistorhinus sepulcralis 12A+A+Y bchrader i960, j brazil it l-iecistor hinus panamensis Kuckes 12A+A+Y Schraaer i960, 1946 Panama 1it Moncus obscurus Dallas 12A+A+Y bchrader 1960a, Panama ’f n Mormidea lumens St§l 12A+A+Y Montgomery 1901, ] 1906, U.S.A. j it Murgantia histrionica btRl 12a +a +Y t Hubhes-ochrader an4i bchrader 1956,USA j it Neodine macraspis Perty 12a +a +Y ochrader 19^ 6, 196( Costa Kica ti Neotti^lossa leporina Futor 12a +a +Y ^chrader 19 6^, Costa i^ica it Neottiglossa pusilla 12a +a +Y bchrader i960, Jakovlev Uosta xxica j t Nezara antennate r^rn- 1£a +a +Y niyamoto 1957, Japan j i University of Ghana http://ugspace.ug.edu.gh AF£tiNi)IX TABLE 10 (cont'd) 121 FAMILY SPECIES 2n REFERENCE AND LOCALITY OF MATERIAL* ^entatominae Nezara viridula St§l 12A+X+Y =Yosida 1956,Japan i f Pentatoma smaragula 12A+X+Y Wilson 1906, U*S.A (Amyotet serville) i i Niphe subferruginea Distant 14A+X+Y Parshad 1957a, India i i Qchrophara montana (List*) 12A+X+Y i-ianna 1951» India i f Palomena angulosa (i'-iontschusky) 14A+X+Y Yosida 1956,Japan i f Palomena prasina Saunders 14a+a+Y Schachow 1932, Germany i i Palomena reuteri (Dist.) 12A+A+Y Parshad 1957b,India i i Palomena viridissima (Poda) i4a+x+y Schachow 1932, Germany i t Pellaea stictica LerK. 12a+a+Y Hughes-Schraaer and schrader 1957*U_Sa i i Pentatoma japonica (i^ ist*) 12A+A+Y Parshad 1957a, India i t Pentatoma rufipes Brulle 12A+A+Y Parshaa, 1957a, Japan t t Peromatus notatus (Amyotet serville) 12A+A+Y Schrader 1945a, Costa x^ ica i i Piezodorus lituratus (Fab.) 12a+^+Y Schachow 1932, i t t t Piezodorus rubrofasciatus(i‘'c Placosternum urus b*)l2A+A+Y 12A+A+Y Germany Manna 1951, India Parshad 1957a,India i i Platycarenus notulatus 12a+a+Y Schrader 1946b, Costa Kica i i f 1 ! Plantia frinbriata j ; j 12a+a+Y Manna, 1958 India j University of Ghana http://ugspace.ug.edu.gh APPENDIX TABLE 10 ( c o n t 'd ) FAMILY SPECIES 2n reference and locality of material* Pentatominae Priossus exemptus (Stal) Pseudovoplitus longicornis (Kuckes) Rhytidolomia saucia Rhytidolomia senilis (Pentatoma) Schraderia cinctus (Kuckes Schraderia hugheseae(Rucker Sciocoris curritans Sciocoris helferii Fieber Sciocoris sulcatus Fieber Scotinophora sp. Scotinophora sp. A Scotinophora sp. £ Scotinophora horvathi(Dist Solubea pugnax St§l Spermatodes sp. Stagonomus bipunctata Linneus Staria lunata Put on Stenozy^um colaratum Klug 12A+X+Y 12A+A+Y 12A+X+Y 14a +a +Y 12a +a +Y )12a +a +Y 12A+X+Y 12A+A+Y 12a +a +Y 10a +a +Y 12A+X+Y 12a +a +Y ) 12A+A+Y 8a +X+Y 14a +a +Y 10a +a +Y 12A+X+Y 12A+A+Y Parshad, 1957 India Schrader 1960b, Costa Rica Schrader I960, U.S.i Schrader I960 Schrader I960, Costa Rica Schrader I960, Costa xiica Parshad 1957i India Jande 1959b, India Miyamoto 1957, Japat Jande 1959b, jLndia Jande 1959b, India Jande 1959b, lnaia rJande 1959b, India i Wilson 1906( Manna, 19511 fcndia Miyamoto 1957, Japan Thyanta antiquensis (Westwcod) 12A+a+Y Thyanta calceata (Say) Thyanta castator (Fab.) 24a +a +Y 14a +a +Y Miyamoto 1957,Japan j Leston and wahrman, j Unpbl. Xsrael j schrader and Iiu^hes- bchrader 1956, Haitij wilson 1906, i ochrader and hu0hes Schrader 1956, USa S chrader and Hu0hes- Sciirader 1956, USA University of Ghana http://ugspace.ug.edu.gh APPJSNDIX TiiBLfl 10 ( c o n t 'd ) f am il y P entat ominae Subfamily Phyloc ephalina< Subfamily Amyoteinae Pentatomidae Amyoteinae i t i i ii SPECIES Thyanta pallidovirens (stal) l^A+A+Y Thyanta pallidovirens (btal) 12a +a +Y Thy ant a perditor (*'ab . ) Thyanta pseudocasta 'Trichopepla semivittata (Say) Podops inuncta (jj'ab.) Arma elector (iJ'abr.) Oechalia grisea (Burm.) Oechalia pacifica otSl Uechalia patruelis ot8l Perillus bioculatus (hineus) 2n 12A+A+Y 12A+A+Y 12a +a +Y 12A+A+Y 12a +a +Y 12a +a +i 12a+a+Y 10a+a+Y 12a +a +Y REFERENCE AND LOCALITY OF MATERIAL * Schrader and Hughes Schrader 1956, USa Schrader and Hughes Schrader I946tlk>A Ueshima 1963 >