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© 2018 The Japan Mendel Society Cytologia 83(2): 165–167

Intraspecific Assessment of philippense Linn. from Uttar Pradesh,

Shashank Kumar Singh and Savariraj Dominic Rajkumar*

Department of Botany, St. Andrew’s College, Gorakhpur, Uttar Pradesh, India

Received November 11, 2017; accepted January 22, 2018

Summary Adiantum philippense Linn. (Adiantaceae) is widely distributed in tropical, sub-tropical and warm temperate regions. Mostly apogamous triploid cytotype (2n=90) has been reported frequently from several lo- calities in India. In the recent exploration of A. philippense species complex in Uttar Pradesh, India, the diploid (n=30) and tetraploid (n=60) cytotypes have been found. The morphological characters such as the lamina, pin- na, spore size, and stoma were measured and compared. The clear difference was detected in spore size between the two cytotypes.

Key words Intraspecific variation, Adiantum, Adiantaceae, Cytotype, Diploid, Tetraploid.

Adiantum philippense Linn. belongs to the family Results and discussion Adiantaceae and it is well distributed in tropical, sub- tropical and warm temperate regions. In India about 25 A. philippense is a common species, widely dis- species of Adiantum have been recorded (Dixit 1984, tributed in the state of Uttar Pradesh and other lo- Chandra 2000, Borthakur et al. 2001). Recently, Fraser- calities of India. The cytological observation revealed Jenkins (2016) has listed 27 taxa of Adiantum of which the presence of a sexual diploid (n=30, Fig. 1E) and eight taxa are exotic, naturalized adventives or widely tetraploid (n=60, Fig. 1G), thereby enlarging the cy- cultivated. About 10 species are reported to be present tological complexity of the species. of diploid in South India (Dixit 1984, Manickam and Irudayaraj were collected in Kushmi Jungle, Gorakhpur, at altitude 1992). A. philippense is a very commonly species found of 84 m (voucher number=SACH-121, 1015), Kushmi in Uttar Pradesh, India. The study area in Uttar Pradesh Jungle, Gorakhpur, 84 m (SACH-129, 161), Pharenda, is bounded by Nepal on the north, states of Uttarakhand Maharajganj, 84 m (SACH-303, 309), Tikari Jungle, on the northeast, Himachal Pradesh on the northwest Gonda, 84 m (SACH-499, 481), Kushmi Jungle, Go- and Bihar on the east. A. philippense is known as a rakhpur, 84 m (SACH-861, 869). Plants of tetraploid complex cytotypes (Verma and Fraser-Jenkins 2008, Bir were collected in Lakhimpur Kheri (Dudhwa National and Verma 2010). In the present exploration from Uttar park), 90 m (SACH-1005, 1009). Kushmi Jungle, Go- Pradesh, India the diploid and tetraploid cytotypes of A. rakhpur, 84 m (SACH-201, 223), Pharenda, Maharajganj, philippense were collected and the intraspecific variation 84 m (SACH-333, 397), Tikari Jungle, Gonda, 84 m between the cytotypes was brought out. (SACH-400, 404), Kushmi Jungle, Gorakhpur, 84 m (SACH-555, 579). Materials and methods In former reports A. philippense known as A. lu- nulatum Burm.f. from Mussoorie and Dehradun (West For cytological study using an acetocarmine Himalaya) reported as an apogamous species (Mehra squash technique (Manton 1950), leaves having young 1932, 1937, 1944) with chromosome count 2n=88 sporangia were collected from the field of and ob- (Mehra 1944). The chromosome count is close to sev- served their chromosome number using a conventional eral subsequent reports of n=90. Roy and Sinha (1961b, technique. Morphological analyses were carried on 1962) had reported 2n=90 (3x, apogamous) in Parasnath characters of lamina, pinna, spore, and sporangia. The Hills, Bihar as well as Assam and Darjeeling (Roy and herbarium specimens were prepared and deposited at St. Sinha 1958, 1961a, b). Somatic chromosome count of Andrew’s College Herbarium (SACH), Gorakhpur. a cultivated 2n=120 (4x, sexual) has also been re- ported in Parasnath Hills (Roy and Sinha 1958). Ghatak (1960, 1963) reported apogamy with 2n=90 (3x, apoga- mous) from West Bengal. Similarly Mehra and Verma (1960, 1963a, b, c), Verma and Loyal (1960), and Roy * Corresponding author, e-mail: [email protected] and Sinha (1961a, b, 1962) from different localities of DOI: 10.1508/cytologia.83.165 India reported mainly triploid apogamous cytotypes. A 166 S. K. Singh and S. D. Rajkumar Cytologia 83(2)

Fig. 1. Herbarium specimen and plant of habitat of diploid (A, B) and tetraploid (C, D) of A. philippense. Meiotic chromosomes and sporangium of diploid (E, F) and tetraploid (G, H), and spores and stomata of diploid (I, J) and tereraploid (K, L). Scale bar=10 µm in E and G and the bar=0.4 mm in F, H–L.

Table 1. Morphological features of the diploid and tetraploid cytotypes of Adiantum philippense.

Measurements Mean value Variance Standard deviation Morphological features 2x 4x 2x 4x 2x 4x 2x 4x

Rhizome Short creeping Short creeping Lamina size (cm) 12–6×7–4 20–13×11–6 8.2×5 15.2×8 4.36×0.8 4.6×2.2 2.08×0.89 2.0×1.48 Total number of pairs 8–12 8–14 9.7 10.8 2.41 3.56 1.55 1.88 of pinnae Pinna size (cm) 2.5–.5×1–.5 3–.5×1.5–.5 1.15×0.82 1.97×1.05 0.3×0.24 0.52×0.48 0.54×0.15 0.721×0.219 Spore size (µm) 27–21×34–29 39–32×45–40 24.3×31.9 35.8×42.9 3.41×2.49 4.73×1.69 1.84×1.57 2.17×1.3 Stoma size (µm) 35.9–30.6×22.7–19.3 44.4–39.3×23.3–20.7 33.03×21.15 42.7×22.4 2.75×1.03 2.72×0.57 1.65×1.01 1.64×0.75 Guard cell size (µm) 33.5–29.2×11.5–9.3 39.7–35.3×14.3–10.4 31.9×10.5 37.5×13 1.85×0.56 1.96×1.46 1.36×0.74 1.40×1.20 tetraploid count of n=60 (4x, sexual) and a morphologi- mean values, variance and standard deviations from ten cally distinct sexual diploid with n=30 in the complex samples each of two cytotypes (Table 1, Fig. 1). The in- were reported from Darjeeling Himalaya by Verma and traspecific cytological complexity has been identified by Mehra (1961) and Mehra and Verma (1963b). Diploid Verma and Mehra (1961) and Mehra and Verma (1963a, apogamous cytotype (n=60) was reported from W. b, c), who discovered four cytotypes in the Darjeeling Himalaya (Khullar and Mehra 1972). Abraham et al. area and the Eastern Indo-Himalaya (Verma and Fraser- (1962) revealed the presence of three cytotypes n=60, Jenkins 2008). There are clear distinction in lamina 2n=120 (4x, sexual) in Ponmudi Hills, 2n=90 (3x, size, stoma size and spore size (Table 1) but on basis of apogamous) in Kerala, and n=30, 2n=60 (2x, sexual) in these differences the cytotypes cannot be separated as Trivandrum. The sexual diploid and tetraploid cytotypes different cytotypes. For intraspecific delimitation also are rare in distribution (Roy and Sinha1958, 1961a, b, some of variation should be present. There seems to be Ghatak 1963, Mehra and Verma 1963b, c, Khare 1980, clear difference in the spore size between the cytotypes Irudayaraj 1998, Bir and Irudayaraj 2001). (Table 1). In Cystopteris the spore size was observed to The present study was done to bring out the intra- increase with the increase in the ploidy level (Hagenah specific complexity of A. philippense complex. The 1961). Wagner (1966) has reported that there were two intraspecific variations have been brought out by mea- varieties of spore size in Gymnocarpium dryopteris, a surement of several characters and calculating their large spore typical in tetraploid and a smaller spore typi- 2018 Intraspecific Assessment of Adiantum philippense from India 167 cal in diploid. In our data the stoma size of the tetraploid distribution of Cystopteris with non-spiny spores in North Amer- cytotype was found to be larger than that of the diploid ica. Rhodora 63: 181–193. Irudayaraj, V. 1998. In: Stace, C. A. (ed.). IOPB Chromosome Data cytotype (Fig. 1L, J). Irudayaraj et al. (1995) have 13. IOPB Newslet. 29: 22–23. observed a variation of spore size between the diploid Irudayaraj, V., Rajkumar, S. D. and Manickam, V. S. 1995 Studies on and tetraploid cytotypes and similar stoma size in both intraspecific variation in South Indian : Rediscovery of the cytotypes in Christella parasitica. All the cytotypes rare diploid cytotype of Christella parasitica (Thelypteridaceae: were found to be sexual with their sporangia including Pteridophyta). Fern Gaz. 15: 41–50. Khare, P. B. 1980. A note on the chromosome number of the fern flora 64 spores. The spores and sporangia were found to be of Amarkantak Hills, Central India. Sci. Cult. 46: 138–139. normal and without any irregularities in their structure. Khullar, S. P. and Mehra, P. N. 1972. Cytotaxonomy of Western Hi- Based on the clear distinction in lamina size, stoma malayan ferns I. Schizaeaceous series. Res. Bull. Sci. Punjab size and spore size in A. philippense intraspecific rank Univ. 23: 186–204. the cytotypes can be assigned because the triploid apog- Manickam, V. S. and Irudayaraj, V. 1992. Flora of the Western Ghats, South India. B. I. Publication Ltd., New Delhi. amous cytotype (2n=90) has not been collected from Manton, I. 1950. Problems of Cytology and Evolution in the Pteri- the present studied area. A cytogenetical survey would dophyta. Cambridge University Press, Cambridge. help to find the triploid apogamous cytotype in other lo- Mehra, P. N. 1932. Occurrence of tracheids in the gametophyte of calities from India to assign the intraspecific rank. Based Adiantum lunulatum. Curr. Sci. 1: 40. on the already existing triploid counts in other parts of Mehra, P. N. 1937. Cytological investigation in the apogamous life India the intraspecific variations could be assessed but cycle of Adiantum lunulatum Burm. Proc. Ind. Sci. Congr. Part III. 21: 266. more authenticity will be achieved only when all the Mehra, P. N. 1944. Cytological investigation of apogamy in Adiantum three cytotypes are collected together and studied. In lunulatum Burm. Proc. Natl. Sci. Inst. Ind. 14B: 189–204. conclusion, the species A. philippense is a puzzling chro- Mehra, P. N. and Khullar, S. P. 1977. Biosystematics of the Adiantum mosomal as well as a reproductive complex which offers lunulatum complex in India with special reference to W. Himala- a challenge to resolve taxonomic issues, despite some yan taxa. Cytologia 42: 501–511. Mehra, P. N. and Verma, S. C. 1960. Cytotaxonomical observa- head-way made by Verma and Fraser-Jenkins (2008). tions on some western Himalayan . Caryologia 13: 619–650. Acknowledgements Mehra, P. N. and Verma, S. C. 1963a. Cytogenetics of Adiantum lunulatum Burm. (s. l.) I. Intraspecific variation and . The authors are thankful to the Principal, St. An- Proc. 50th Indian Sci. Congr. Part III. Sect. 6. pp. 454–455. Mehra, P. N. and Verma, S. C. 1963b. Cytogenetics of Adiantum lu- ’ drew s college (PG), Gorakhpur, Uttar Pradesh, India for nulatum Burm. (s. l.) II. Cytogenetic interpretation of intraspecif- the facilities and the encouragement given to us. ic variation. Proc. 50th Indian Sci. Congr. Part III. Sect. 6. p. 454. Mehra, P. N. and Verma, S. C. 1963c. Polymorphicity and cytoge- References netics of Adiantum lunulatum complex. J. Ind. Bot. Soc. 42A: 110–121. Abraham, A., Ninan, C. A. and Mathew, P. M. 1962. Studies on the Roy, R. P. and Sinha, B. M. B. 1958. Somatic studies in the genus cytology and phylogeny of the VII. Observations Adiantum. Proc. 45th Indian Sci. Congr. Part III, Sect. 6. pp on one hundred species of south Indian ferns. J. Ind. Bot. Soc. 284–285. 41: 339–421. Roy, R. P. and Sinha, B. M. B. 1961a. Meiotic studies in the genus Bir, S. S. and Irudayaraj, V. 2001. Cytology of some ferns from the Adiantum. Proc. 48th Indian Sci. Congr. Part III, Sect. 6. pp. Nilgiris, South India-IV. Fern Gaz. 16: 177–190. 264–265. Bir, S. S. and Verma, S. C. 2010. Chromosome Atlas of the Indian Roy, R. P. and Sinha, B. M. B. 1961b. Meiotic studies in the genus Pteridophytes (1951–2009). Bishen Singh Mahendra Pal Singh, Adiantum. Caryologia 14: 413–428. Dehra Dun. Roy, R. P. and Sinha, B. M. B. 1962.Cytotaxonomic Studies in the Borthakur, S. K., Deka, P. and Nath, K. K. 2001. Illustrated Manual of Genus Adiantum. In: Maheshwari, P., Johri, B. M. and Vasil, I. Ferns of Assam. Bishen Singh Mahendra Pal Singh, Dehradun. K. (eds.). Proc. Sum. Scho. Bot., Darjeeling, CSIR, New Delhi. Chandra, S. 2000. The Ferns of India, Enumeration, Synonyms and pp. 349–362. Distribution. International Book Distributors, Dehradun. Verma, S. C. and Fraser-Jenkins, C. R. 2008. Adiantum philippense Dixit, R. D. 1984. A Census of Indian Pteridophytes, Flora of India, L. the Correct Name for A. lunulatum Burm. F. and Its Subspe- Series IV. Bot. Surv. India, Howrah. cies. Perspectives in Pteridophytes, Bishen Singh Mahendra Pal Fraser-Jenkins, C. R. 2016. A revised checklist of Indian pterido- Singh, Dehradun. pp. 65–92. phytes-I. Ind. Fern J. 33: 193–205. Verma, S. C. and Loyal, D. S. 1960. Chromosome counts in some Ghatak, J. 1960. Apogamy in Adiantum philippense L. and its cytol- ferns from Nainital. Curr. Sci. 29: 69–70. ogy. Proc. 47th Indian Sci. Congr. Part III, Sect. 6. pp. 341–342. Verma, S. C. and Mehra, P. N. 1961. Chromosome numbers in Hima- Ghatak, J. 1963. Apogamy in Adiantum philippense L. and its cytol- layan ferns. Res. Bull. Punjab Univ. Sci. 12: 139–164. ogy. Bull. Bot. Soc. Bengal 13: 63–65. Wagner, W. H. Jr. 1966. New data on North American oak ferns, Hagenah, D. J. 1961. Spore studies in the genus Cystopteris L. The Gymnocarpium. Rhodora 68: 121–138.