OCCASIONAL PAPER NO. 300 Records of the Zoological Survey of India Part-l : A review of studies on the chromosomes of Grasshoppers in India (1928-2006) ASHOK K. SINGH Part-2 : The nuclear phenotype of Xenocatantops humilis (Serville) (Orthoptera : Acrididae : Catantopinae), w.s.r. to supernumerary segments ASHOK K. SINGH Part-3 : Supernumerary chromosomes of Patanga succincta (Johansson) (Orthoptera : Acrididae : Cyrtacanthacridinae) ASHOK K. SINGH V. B. SINGH ZOOLOGICAL SURVEY OF INDIA OCCASIONAL PAPER No. 300 RECORDS OF THE ZOOLOGICAL SURVEY OF INDIA Part·1 : A Review of studies on the chromosomes of Grasshoppers in India (1928·2006) Part·2 : The nuclear phenotype of Xenocatantops humilis (Serville) (Orthoptera : Acrididae : Catantopinae), w.s.r. to supernumerary segments ASHOK K. SINGH Cytotaxono,,,y Research Laboratory, Zoological Survey of India 27, lawahar Lal Nehru Road, Kolkata- 700016 Part·3 : Supernumerary chromosomes of Patanga succincta (Johansson) (Ortboptera : Acrididae : Cyrtacanthacridinae) ASHOK K. SINGH Zoological Survey of India, 27, lawahar Lal Nehru Road, Kolkata-700016 and V. B. SINGH Departmel1l of Zoology, Uda, Pratap Autonolnous College, Varanasi- 221002 Edited by the Director, Zoological Survey of India, Kolkata Zoological Survey of India Kolkata CITATION Singh, 2009. A Review of studies on the chromosomes of Grasshoppers in India (1928- 2006) (Part-I) : 1-41 and The nuclear phenotype of Xenocatuntops "utlli/is (Serville) (Orthoptera : Acrididae : Catantopinae), w.s.r. to supernumerary segments (Part-2) : 43- 61; Singh and Si~gh, 2009. Supernumerary chromosomes of Patanga succincta (Johansson) (Orthoptera : Acrididae : CYl1acanthacridinae) (Part-3) : 63-72. Rec. zool. Surv. India, Occ. Paper No., 300 (Published by the Director, 2001. Surv. India, Kolkata) Published : August, 2009 ISBN 978-81-8171-227-1 © Govt. of India, 2009 All RIGHTS RESERVED • No part of this publication may be reproduced, stored in a retrieval system or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise without the prior permission of the publisher. • This book is sold subject to the condition that it shall not, by way of trade, be lent, re-sold, hired out or otherwise disposed of without the publisher's consent, in any form of binding or cover other than that in which it is published. • The correct price of this publication is the price printed on this page. Any revised price indicated by a rubber stamp or by a sticker or by any other means is incorrect and should be unacceptable. PRICE India : Rs. 150.00 Foreign: $10; £ 7 Published at the Publication Division by the Director, Zoological Survey of India, 234/4, A.J .C. Bose Road, 2nd MSO Building, Kolkata-700 020 and printed at Calcutta Repro Graphics, Kolkata-700 006 Records of the Zoological Survey of India OCCASIONAL PAPER Mo.. 2009 1-72 CONTENTS Part-l : A Review of studies on the chromosomes of Grasshoppers in India (1928-2006)..................................................................................................... 1-41 Part-2 : The nuclear phenotype of Xenocatantops humilis (Serville) (Orthoptera : Acrididae : Catantopinae), w.s.r. to supernumerary segments .......... 43-61 Part-3 : Supernumerary chromosomes of Patanga succincta (Johansson) (Onhoptera : Acrididae : Cyrtacanthacridinae) ......................................................... 63-72 Zoo I. Surv. I"dia Rec. zool. Surv. India, Occ. Paper No. 300(Part-l) : 1-41,2009 A REVIEW OF STUDIES ON THE CHROMOSOMES OF GRASSHOPPERS IN INDIA (1928-2006) ASHOK K. SINGH Cytotaxonon,y Research Laboratory Zoological Survey of India 27, lawahar Lal Nehru Road, Kolkata-7000J6 INTRODUCTION Three hundred ninety-three publications (including 200 abstract papers) have been recorded on Indian grasshoppers. 117 species of the grasshoppers have been utilized which revealed prolific choice of this insect as study material. Grasshoppers have also been extensively used in screening mutagenic and carcinogenic potential of a variety of chemicals and drugs (for total information please refer publication viz., Singh, A. K. 2002. A Catalogue of Experimental Studies on Grasshopper Chrolnosol1zes ill India 1928 to 2000. Rec. zoof. Surv. India, Occ. Paper No. 197 : 1-44.) The present review is a subject-wise presentation of the findings on the chromosomes. GENERAL CYTOGENETICS A Karyology Karyological characterization of grasshopper species for cytotaxonomical study, and detection of gross aberrations in chromosome number, morphology and sex-chromosome mechanism were the objectives outlined by the earlier cytologists. The group has long been recognized as chromosomally conservative. Majority of species have 19 xoa; 20 XX ~ (Pyrgomorphidae) and 23 xoa; 24 XX ~ (Acrididae) with all the members as rods. A variety of deviations from this uniformity have however been recorded in literature (Whitel973; Hewitt 1979). A serious problem encountered in karyology is genetically controlled variation in contraction within the chromosome complement of a cell. In addition, an "artificial" variation can also be introduced by the experimenter viz., on excessive concentration and length of pretreatment 2 Rec. zool. Surv. India, Dcc. Paper No. 300(part-l) with colchicine leading to differential contraction of euchromatin and heterochromatin (Smith 1965). On reliability of banding methods now in extensive use in karyology, utility of correct determination of chromosome length and arm length have been reduced. Underlying gross structural heterozygosities are also detected considerably. Study of the karyological features of a species and its populations are essential for understanding the evolution of the chromosomal changes in them. i) Chromosome number and morphology Cytqgenetic investigations on animal chromosomes in India started as early as 1928. It was Asana who initiated it with his observations on the spermatogenesis of a grasshopper species viz., Poeeiloeera pieta. The species has 2n = 19 with XO sex chromosomes in males. In 1931 Asana studied the spermatogenesis of 6 species belongi ng to the 4 subfamilies viz., Tryxalinae, Oedipodinae, Catantopinae and Pyrgomorphinae. He and his associate Mahabale in 1945 reported spermatogenesis of the desert locust Sehistoeerea gregaria. Asana in collaboration with Makino (1934) described the spermatogonial metaphase chromosomes of Poecilceerus pietus. With a cytotaxonomical approach Asana et ale (1939) analysed the chromosome complements of 6 species belonging to the subfamilies Acridinae, Oedipodinae and Catantopinae. They found that small dot like chromosomes were in no way characteristic of a subfamily. One, two or three such pairs were although recorded, but not uniformly in all the species studied under a subfamily. Rao made fairly extensive studies during 1932 to 1940 to trace chromosomal relationships in Indian Pyrgomorphinae. His comparative account of the chromosomes of eight genera of Indian Pyrgomorphinae (Rao 1937) is considered as the first comprehensive study of cytotaxonomical nature in India. He discovered that the size of chromosomes varied considerably in different genera. However, on analyzing the measurements of meiotic metaphase II chromosomes for sizes and seriation, there was some degree of uniformity within a subfamily. In this study he has shown that simple elimination of two pairs from the usual twenty three chromosomes of the complement in Acrididae does not explain fully karyologically any phylogenetic relationship of Pyrgomorphinae with Acrididae. Ray-Chaudhuri along with his collaborators started cytogenetic studies on grasshoppers in the late 1940s and made a substantial contribution particularly in the field of radiation cytogenetics. He ha~ first started research in Drosophila genetics at the Zoology Department, Calcutta University. But due to several technical limitations he switched over to work on the chromosomes of grasshoppers and other arthropods. Later he had revived his interest in Drosophila genetics with the help of Prof. 1.B.S. Haldane who had reestablished Drosophila genetics laboratory for him in the Department of Zoology, Calcutta University. By this time research laboratories in universities tackled a variety of problems of fundamental importance. Srivastava in Allahabad, Sharma in Chandigarh, Manna in Kalyani, Rajasekarasetty in Mysore~ Sharma, T in Banaras and Yadav in Kurukshetra, have been the major contributors in this field. SINGH: A rrvi,w of studies on tire clrronlosonles of grasshoppers in India (/928-2006) 3 Ray-Chaudhury and Dutt (1947) presented a comparative study of the chromosome structure and behaviour in three different genera, viz., SpathosternUl1l, Oxya and Phloeoba. They showed that an indirect proportionality between length and number of chiasmata existed in all the s~cies studied, and terminalization coefficients failed to show any significant increase from diplotene to metaphase I. They reported presence of a chiasma in the short arms or'the ring bivalents of Oxya sp. There is however controversy existing regarding occurrence of chiasma in the short arms of acrocentric chromosomes. Dult (1948a) on cytological investigation of genus Phlaeoba, recognized two different forms and assigned them as forms A and B. He sent the specimens to B.P. Uvarov of Imperial Bureau of Entomology in London for identification, who identified them as two separate species but had not assigned specific names to them. However, B.R. Seshachar of Bangalore identified them as one species Phlaeoba angusfidorsis.