_??_1988 by Cytologia, Tokyo C ytologia 53: 679-684, 1988

Cytotaxonomical Studies of South Indian Capparidaceae

D. Subramania and G. Susheela

Department of Botany, Annamalai University , Annamalainagar-608002, Tamilnadu ,

Accepted October 9, 1987

The capparideceae consist of about 46 genera and nearly 700 species (Lawrence 1957) . Willis (1966) mentioned 30 genera and 650 species . There are 7 genera including 32 species of capparidaceae described by Gamble (1965) in his "Flora of presidency of Madras" occurring in Madras and neighbouring states , Andhrapradesh and Karnataka. Cytological studies in South Indian species of Capparidaceae are fragmentary and poor and therefore , the present investigation has been undertaken in 18 species coming under 6 genera .

Materials and methods

Most of the species were locally obtained . Cleome felina was collected at courtallum of . Maerua arenaria, Gapparis brevispina and C. sepiaria were obtained from Christian College campus, Tambaram, Madras. C. divaricata was collected from Maruthamalai of Coimbatore District and Cadaba triphylla from Mundanthurai of Tirunelveli District. Seedlings of the

species were grown in our Botanic Garden, Annamalai University in 1•Œclay pots . For mitotic studies, healthy root tips were pretreated in 0.02% hydroxyquinoline kept at 4•Ž for 3hours. Then, they were washed in distilled water and fixed in 1: 3 acetic ethanol For at least one hour or overnight. Following the method of iron alum haematoxylin squash technique described by Marimuthu and Subramaniam (1960), root tip squashes were made and the slides sealed. Important plates were photographed and camera lucida diagrams were Irawn with Abbe camera lucida under oil immersion lens at a magnification of•~1250.

9 plates were considered for karyotype analysis, in each species studied. The measurements of chromosomes were made with ocular micrometer, the scale of which had been calibrated, from stage micrometer. From the absolute chromosome lengths of the species studied, histogram was drawn.

Observations

The chromosomes of the members of the family Capparidaceae are generally short (Figs. I to 7). Based on the length, the chromsomes are grouped into shorter, short, medium and long

types (Battaglia 1955). Under each of these groups, the following categories of chromosomes have been recognised. Long (Above 2.5 to 3.0ƒÊm)

Type A: Chromosome with submedian primary centromere and subterminal secondary cen tromere. Type B: Chromosome with median primary centromere and subterminal secondary cen tromere. Type C: Chromosome with submedian centromere.

Type D: Chromosome with median centromere. 680 D. Subramanian and G. Susheela Cytologia 53

Figs. 1 to 7: Somatic chromosomes. 1, Cleome viscosa, 2n=20. 2, C. monophylla, 2n=22. 3, Cadaba indica, 2n=34. 4, Cleome chellidonii. 2n=34. 5, diversifolia, 2n=98. 6, C. grandis, 2n=42. 7, Crataeva religiosa, 2n=26.

Table 1. Particulars about the present and previous reports of chromosome numbers (vide Fedorov 1974 and Taxon reports upto 1983) 1988 Cytotaxonomical Studies of S outh Indian Capparidaceae 681

Medium (Above 2 .0 to 2.5ƒÊm). Type E: Chromosome with sub-median primary centrom ere and subterminal secondary centromere. Type F: Chromosome with submedian centromere . Type G: Chromosome with median centromere .

FIG-8

Fig. 8. Histogram. Fig. 9. Graph.

Table 2.

The descriptions of the various columns of Table 2 are as follows: 1, Chromosome number (2n). 2,

size range in millimicrons. 3, Number of chromosomes with secondary constrictions. 4, Number of chro mosomes with submedian centromere. 5, Number of chromosomes with median centromere. 6, Number

of chromosomes with subterminal centromere. 7, Absolute chromosome length (Fig. 8) in ƒÊm. 8, Average chromosome length in ƒÊm. 682 D. Subramanian and G. Susheela Cytologia 53

Figs. 10 to 36. Drawings of meiosis. 10 to 12. Gynandropsis pentaphylla. 10, I prophase. 11, Diakinesis. 12, I metaphase, n=16. 13 to 16, Cleome viscosa. 13, I metaphase with per sistent nucleolus. 14. I metaphase, n=10. 15, II metaphase, polar view. 16, II metaphase side view with precocious movement of chromosome. 17 to 19: Cleome monophylla. 17, I metaphase, n=11. 18, I metaphse with 10 bivalents, and 2 univalents. 19, I metaphase with 22univalents. 20 and 21, Cleome tenella. 20. I metaphase (n=9). 21, I metaphase (n=10). 22 to 25: Cleome felina. 22, I metaphase (n=10) with persistent nucleolus. 23, II metaphpase, polar view. 24 and 25, I anaphasic laggards and bridges. 26 to 28: Cleome aspera. 26, I metaphase with quad rivalents and 5 bivalents. 27, I metaphase with 2 quadrivalents and 9 bivalents. 28, anaphasic laggard. 29 and 30 Cleome chellidonii. 29, I metaphase with 13 bivalents and 2 quadrivalents. 30. II metaphase. 31 and 32: Cadaba indica. 31 and 32. I metaphase n=17. 32=I1 met aphase. 33. Cadaba triphylla, I metaphase, n=17. 34 and 35, Capparis sepiaria. 34 aphase. . 35, tetrad formation showing the chromatins. 36, Capparis zeylanica, I metaphase,I met n=22. 1988 Cytotaxonomical Studies of South Indian Capparidaceae 683

Short (Above 1.0 to 2.0ƒÊm). Type H: Chromosome with median primary centromere and subterminal secondary cen

tromere. Type I: Chromosome with submedian centromere.

Type J: Chromosome with median centromere .

Type K: Chromosome with subterminal centromere . Shorter (From 0.6 to 10ƒÊm)

Type L: Chromosome with submedian centromere . Type M: Chromosome with median centromere. Type N: Chromosome with subterminal centromere. The total number of secondary constricted chromosomes has been found to vary from species to species. The karyomorphological features of the various species studied are sum

marised in Tables 1 and 2. Meiotic studies have been made in Gynandropsis pentaphylla, Cleome viscosa, C. mono

phylla, C. tenella, C. felina, C. aspera, C. chellidonii, Cadaba indica, C. triphylla, Capparis sepiaria and C. zeylanica (Figs. 10 to 36). In the species in which both mitotic and meiotic studies have been made the meiotic chromosome numbers are in correlation with the respective mitotic chromosome numbers.

Discussion

First record of chromosome number has been made in Cleome tenella, C. felina, C. aspera,

Cadaba triphylla, Capparis brevispina, C. diversifolia, C. rotundifolia and C. divaricata (Table 1.) Deviant report as against the previous record of chromosome number has been made in Maerua arenaria, Capparis zeylanica and Cadaba indica (Table 2). In all other species studied, the present report of chromosome numbers are in confirmation with the earlier reports. The basic chromosome number ranges from n=9 to n=80. The graph drawn on the basis of frequency distribution of basic chromosome numbers shows a polymodel curve (Fig.

9) showing thereby the evolution of species in multivarious lines and indicating the polyphyletic nature of the species. This is sufficiently evidenced and explained by the earlier contributions of Raghavan (1938) and Raghavan and Venkatasubban (1939 and 1940). According to them, the evolution of the genera seemed to have followed irregular lines.

Secondary association of the bivalents had been observed by Raghavan and Venkatasub ban (1939) and the basic number based upon maximum association was 7. In the present investigation the basic number of chromosomes is 9 in Cleome tenella and 10 in C. viscosa and C. felina. These basic numbers, along with n=11 and n=13 might have arisen from the prima ry basic number n=7 by aneuploidy. The basic numbers n=16, n=17 and n=18 are secon

darily balanced tetraploids derived from n=7 by means of both euploidy and aneuploidy. The basic numbers n=20, n=21 and n=22 may be secondarily balanced higher polyploids. The basic numbers n=49 and n=80 may be secondarily balanced allopolyploids which

might have been evolved by hybridization of any two species followed by polyploidization. More or less similar conclusions have been drawn by the cytological investigations in this family by Raghavan and Venkatasubban (1941) and Raman and Kesavan (1963). Therefore,

C. tenella, C. felina, C. viscosa, C. monophylla, C. aspera and Crataeya religiosa are examples for aneuploids. Gynandropsis pentaphylla, Cleome chellidonii, Cadaba indica, C. triphylla and Capparis brevispina are instances for secondarily balanced tetraploids. Maerua arenaria,

Capparis grandis, C. rotundifolia, C. sepiaria and C. zeylanica are examples for secondarily balanced higher polyploids. Capparis diversifolia and C. divaricata are instances for secon darily balanced allopolyploids. Such a possibility of derivation of secondarily balanced al- 684 D. Subramanianand G. Susheela Cytologia53

lopolyploids in the origin and evolution of species has been described among the south Indian species of Acanthaceae (Govindarajan and Subramanian 1983) and Araceae (Ramachandran 1978). The chromosomes of the species studied are generally smaller in size. In this character, the species of Capparidaceae are closely related together and highly evolved.

Summary Cytotaxonomical studies have been made in 18 species coming under 6 genera of South Indian Capparidaceae, of which first record of chromosome number has been made in Cleome tenella, C. felina, C. aspera, Cadaba triphylla, Capparis brevispina, C. diversifolia, C. rotundi folia and C. divaricata. The basic chromosome number ranges from n=9 to n=80. The graph drawn on the frequency distribution of basic chromosome numbers shows a polymodel curve indicating thereby the evolution of species in many directions. The basic chromosome number may be n=7, from which species with n=9, n=10, n=11 and n=13 might have arisen by aneuploidy. The basic numbers n=16, n=17 and n=18 are secondarily balanced tetraploids by means of euploidy and aneuploidy. The basic numbers n=20, n=21 and n=22 may be secondarily balanced higher polyploids. The basic numbers n=49 and n=80 may be secondarily balanced allopolyploids which might have been evolved by hybridization of any two species followed by polyploidization. Therefore, autopolyploidy as well as allopolyploidy might have played important role in the the origin and evolution of the species of South Indian Capparidaceae.

Acknowledgements The authors express their heartfelt thanks to Dr. R. Ganesan, Professor and Head of the Dept. of Botany, Annamalai University, for his encouragements and facilities provided through out the course of this investigation. References Bir, S. S. and Sidhu, Manjit 1974. IOPS chromosomenumber reports. Taxon 23(2/3):373-380. Fedorov,A. N. A. 1974. ChromosomeNumber of FloweringPlants. Reprintsby Otto KoeltsScience Pub lishers,No. 624,Koenigstein, West Germany. Gamble,J. S. 1956. Flora of presidencyof Madras. Vol. II. B. S. I. Publication,Calcutta. Govindarajan,T. and Subramanian,D. 1983. Karyomorphologicalstudies in SouthIndia Acanthaceae. Cyto logia48: 491-504. Janaki Ammal,E. K. 1933. The chromosomenumber of Cleomeviscosa. Curr. Sci. 1: 328. Lawrence,G. H. M. 1957. Taxonomyof VascularPlants. The MacmillanCo., New York. MarimuthuK. M. and Subramaniam,M. K. 1960. A haematoxylinsquash method for the root tips of Dolichos lablabL. Curr. Sci.29: 482-483. Raghavan,T. S. 1938b. Morphologicaland cytologicalstudies in the CapparidaceaeII. Floral morphology and cytologyof Gynandroosispentaphylla Dc. Ann. Bot. 2(5): 75-95.- and Venkatasubban,K. R. 1939. Studiesin the Capparidaceae.V. The cytology of Crataeva religiosa Cytologia10(1-2): 23-31. - and Venkatasubban,K. R. 1941. Studiesin the Capparidaceae.VIII. The cytologyof Cappariszeylanica Linn.and relatedgenera. Cytologia11(3): 319-331. Ramachandran,K. 1978. Cytologicalstudies in South Indian Araceae. Cytologia43: 289-303. Raman, V. S. and Kesavan,P. L. 1963. Chromosomenumbers of some dicotyledons. Sci. and culture 29(8):413-414. Willis,J. C. 1966. Dictionaryof the FloweringPlants and Ferns. CambridgeUniversity Press. London.