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 <ihana, Jbegon; and confirmed at the Commonwealth Institute 
of iioatomology, .London*
University of Ghana          http://ugspace.ug.edu.gh
SECTION 3
RESULTS
3#1 INTRODUCTION
The Heteroptera are supposed to be generally an easy cytological 
material, because of their deep staining chromosomes, and heteropycnotic 
sex chromosomes* Their chromosomes according to Hanna (1951)» “have no 
longitudinal differentiation by which one chromosome can be distinguished 
from the other11, and therefore, in families like Pentatomidae where the 
chromosome number is constant in different species it becomes difficult 
to compare their chromosome complements. The only other convenient 
alternative in such a situation is to analyse tne karyotypes according to 
the size of the chromosomes, and to classify them into different size 
groups* A visual estimation of the size of the spermatogonial and first 
meiotic _metaphase_chromosomes has been the usual method employed in these 
analyses to evaluate phylogenetic relationships.
3.2 MEIOSIS
The general course of meiosis in all the families studied is fairly 
uniform and the morphology and behaviour of the chromosomes of the group 
are typical heteropteran. The diploid number of chromosomes is expressed 
during the spermatogonial metaphase, and tne haploid numbers at the 
spermatocytic metaphase one. In the earliest spermatocyte stage, the 
chromosomes are generally found in a more or less clumped condition at
University of Ghana          http://ugspace.ug.edu.gh
the central area of the nucleus, leaving a clear gap near the nuclear 
membrane* 'Uhe sex chromosomes which are positively heteropycnotic, 
may sometimes be observed to lie within the central chromosomal mass 
and sometimes very close to the nuclear membrane* In the early 
diplotene stage, the tetrad nature of each bivalent becomes evident.
Much condensation of euchromatic portions takes place through diplotene.
Sex chromosomes, if fused, separate at the prometaphase stage and 
lose some of their heteropycnotic character* At this point sex-chromosomes 
can be distinguished from the autosomes, because they are composed of 
only two instead of four chromatids as in autosomal bivalents*
At the first metaphase stage (metaphase I), the autosomal bivalents 
arrange themselves roughly in the form of a ring on the equatorial 
plate, and the central part of the plate is generally occupied by one 
or both the sex chromosomes* when there is only one sex chromosome at 
the centre, the other generally takes up its position along with the 
bivalents* In very few cases, the central position is occupied by an 
autosome* Wormally, hov/ever more than two elements are not present in 
the central region* The different species among the various families 
behave differently in these respects* At the second spermatocytic 
metaphase (metaphase II), the sex chromosome pair almost invariably lie 
at the centre (unseparated) surrounded by a rin& of autosomes* At 
Anaphase II, two types of chromosome distribution result, following 
the segregation of the sex—chromosomes, the X“cto°m0isc>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- ' -
•
- <r------x
f 0
• t
29 3 0
lOju
3.3*7 Figs. 27-30: Aspavia hastator (Fabr.) 2?) microphotograph, 28) tracing 
of Diakinesis, 29) microphotograph, 30) tracing of hetaphase II, polar 
view.
University of Ghana          http://ugspace.ug.edu.gh
Figs* Atelocera serrata (Fabr*) 1^) microphotograph, 32) tracing
of jJiakinesis, 33) microphotograph, J4) tracing of hetaphase II, 
polar view.
University of Ghana          http://ugspace.ug.edu.gh
figs. 35-38: Bathycoelia thalassina i,Ji.-s) : 35) microphotograph, 
36) tracing of Ldakinesis with 7 elements, 37) microphotograph, 
38) tracing of hetaphase II, polar view.
University of Ghana          http://ugspace.ug.edu.gh
3.3»10 iJ'igs. 39-42: Bathycoelia rodhaini (schout.) 39) microphoto^raph,
40) tracing of Metaphase II, polar view with 7 elements,
41) microphotograph, 42) tracing of netaphase II, side view.
University of Ghana          http://ugspace.ug.edu.gh
i p y p f c
s
44
lOA'
11 Figs. 3^-46: Benia sp. a. 3^) microphotograph, **4) tracing (■£ 
microphotograph size) of Diakinesis, microphotograph,
6^) tracing (•£ microphotograph size) of hetaphase I.
University of Ghana          http://ugspace.ug.edu.gh
VI V 1 " '
t  r
50
.
# . *
♦
> ^  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<T)CT)CJl^ hOOCJ> 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)
<T)
to
00
: n
Amyoteinae
r rv<-oooT5ITQ
5*o
Podopinae
Pentatominae
SUB 
FAMILY
—1 -*-* —» ro —*4N 4S ro ro —*• —* —» —» —x -jcn cd cn ^  w O cd
rOZ1zo
CO —* —i. — - —“—k O') —fc o  U> —» -*
”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 <X> 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 ±<uhekerns mit besonderer
Berucksichtigung der Jieteropteran. Biol, abl. .Leipzig 
£8: 152-179.
14. HiiLKiiti, U. 1956. studies on mitotic and meiotic cell division
in certain Hemiptera under normal ana experimental 
conditions. Ann. Acad, oci. 32: 1-80.
15* HElki^ K, P. 1951* The chromosome cytology of an Oahu species of
the Pacific genus uechalia (Pentatomidae, hemiptera- 
iieteroptera). J. Morph, 88: I85-I98.
16. HUviiiiijB-SCiiJ^ADiaL'i, b. and Kl6, d. 1941. The diffuse spinale attach­
ment of coccids, verified by the mitotic behaviour of 
inauced chromosome fragments. J. expt. zool. Philad.
8Z: 429-456.
University of Ghana          http://ugspace.ug.edu.gh
1942. i'he chromosomes of Nautococcus 
bchraderae Vays and the meiotic division 
figure of male Haveiinecoccids. J. horph.
20: 261-300.
bm and SCiiKAi->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<es. Bull. Panjab. Univ.
10 Pt 1: 25-34.
JAi'JUni, b.«b. 1959b. Chromosome number and sex Mechanism in twenty- 
seven species of Inaian heteroptera.
Jtes. -bull. Pan.jab. Univ. 10 Pt XI. 215-217* 
o.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, PMDjCiRGi<AbT, J.6. and BOUTHwOOu, T.R.B, 195^ » Classi­
fication of the terrestrial Heteroptera. Nature 
174: 91-92.
39• *Ii£THII'iRRY et 1&93* Catalogue general des Hemipteres
Musee royal d^istoire naturelle de Belgique 1: 239* 
kOm L0UIB and KUMaR, H. 1972. Morphology of the alimentary and
reproductive organs in Keduviiaae (Hemiptera: 
Heteroptera) with comments on interrelationships 
within the family. Ann. -Edit, aoc. Amer. 66 (3) :635“638.
41. MACKBRUAB, I.i'-x. £ed.) 1970. The Insects of Australia.
Melbourne University Press. 1029 pp.
42. MAKING, B. 1951* An atlas of the chromosome numbers in animals.
Iowa Btate College Press, iimes. Iowa.
University of Ghana          http://ugspace.ug.edu.gh
43# maxima, U.K. ,1950. Multiple sex chromosome mechanism in a
reduviid bug. Conorhinus rubrofasciatus (de Greer). 
Proc. zool. £>oc. 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 > <Jaoan
Schrader and Hughesf- 
Schrader 1956, UsA 
Montgomery 1901, 
1906; Wilson 1906, 
U«o«A»
Kumar (Unpb1.)Japar
woodward 1950; 
Pendergrast 1957, 
England
Miyamoto 1957, 
Japan
Heizer 1951 ,Hav/ai 
Heizer 19511 1 
Heizer 1951, 1
»iilson 1906, U»q «a
University of Ghana          http://ugspace.ug.edu.gh
APPiJM£X TA13LE 10 (c o n t * d )
family
Pentatomidae
^myoteinae
SPECIES
Perillus confluens stal 
Picromerus bidens (Amyot et 
Serville)
Picromerus niffridens Puton 
Podisus bracteatus (Fitch) 
Podisus macuventris (bay)
Podisus modestus (oay) 
Podisus placidus Uhler 
btiretcus anchorafto (Fabr.)
Troilus huridus (Fabr.)
2n
12A+A+Y
12A+a+Y
12a +a +Y
12A+A+Y
12a+a+Y
14a +a +Y
14a +a +Y
12a +a +Y
12a +a +Y
REFERENCE AND 
LOCALITY OF 
MATERIAL *
Montgomery 1901*U s a  
Yosida 1956, Japan
Miyamoto 1957? JapaL
Wilson 1906b,
Montgomery 1901,
U«o»A»
wilson 190bb, U «o»A
Wilson 1906 , i *Q ♦ A<
Parshad 1957a, 
India
Miyamoto 1957* 
Japan
* Locality of 
material under­
lined
University of Ghana          http://ugspace.ug.edu.gh
University of Ghana          http://ugspace.ug.edu.gh