Shekhar Pandey C
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T REPRO N DU LA C The International Journal of Plant Reproductive Biology 8(2) July 2016, pp.202-207 P T I F V O E Y B DOI 10.14787/ijprb.2016 8.2. 202-207 I T O E I L O C G O S I S E T S H Pollen morphology in Inula L. (Asteraceae) and related genera in India T Shweta Shekhar1 and Arun K. Pandey2* 1Department of Botany, H.N.B. Govt. Post-Graduate College, Naini-211008 (Allahabad), India 2Department of Botany, University of Delhi, Delhi-110007, India *e-mail : [email protected] Received : 19.02.2016; Revised: 10.04.2016; Accepted and Published on line: 01.07.2016 ABSTRACT In Inula (s.s.), Duhaldea, Dittrichia and Iphiona pollen grains are usually radially symmetrical, isopolar, prolate-spheroidal rarely oblate-speroidal (Inula obtusifolia), mostly tricolporate, rarely tetracolporate (Iphiona grantioides) and zono-aperturate. Spines are mostly acute-acuminate or sometimes rounded-blunt along with few acute apices (Inula rhizocephala). On the basis of spines apices, P/E ratio and polar length, presently studied taxa are grouped under 3 major types viz., (1) Inula acuminata-type (2) Inula rhizocephala-type and (3) Duhaldea cappa-type. Keywords : Inula, Asteraceae, pollen, Taxonomic Considerations. The genus Inula L. (tribe Inuleae, sub-tribe Inulinae, The concept of the genus Inula (s.l.) is broad and family Asteraceae) includes c. 100 species mainly heterogenous. This has resulted into an assemblage of several distributed in warm and temperate regions of Africa, Asia unrelated taxa under the broad canopy of Inula thus causing lot and Europe (Anderberg 2009, Chen and Anderberg 2011). of taxonomic confusions. In view of taxonomic problems, the In India, the genus Inula (s.l.) has been reported to contain present study on pollen morphology of Inula (s.s.) and allied 20 species (Kumar and Pant 1995) distributed mainly in genera has been undertaken to utilize the data for taxonomic Trans Himalaya, Western Himalaya and North East regions. purposes. Our taxonomic revision on the genus Inula (s.s.) in India MATERIAL AND METHODS (Shekhar et al. 2013) recognizes 12 species; whereas, In the present study 15 taxa belonging to the genus Inula remaining species reported in Flora of India have been (s.s.), Duhaldea, Dittrichia and Iphiona from India has been transferred to the genera Duhaldea, Iphiona and Dittrichia studied from palynological point of view. Pollen grains of following nomenclatural changes made by Anderberg Inula (s.s.) and its allied genera were examined from anthers (1991). collected directly in the field or from herbarium specimens. The study of the pollen morphology in tribe Inuleae Data on pollen characters are given in table 1. Pollen grains (Asteraceae) has been carried out by a number of workers were acetolysed (Erdtman 1969) and viewed using both an (Wodehouse 1926-1935, Stix 1960, Skvarla and Larson Olympus Magnüs microscope (MLM 90 703518) and SEM. 1965, Skvarla and Turner 1966, Singh and Joshi 1969, For SEM studies, pollen grains, passed through Dakshini and Singh 1970, Payne and Skvarla 1970, Lenis dehydration series, were suspended in a drop of acetone and 1971, Keeley and Jones 1977, Skvarla et al. 1977, Bolic then transferred onto a 5 mm round glass coverslip using 1978, Robbins et al. 1979, Clark et al. 1980, Blackmore 1981, Nair and Lawrence 1985, Robinson and Marticoena double-sided adhesive tape. The material was coated with a 1986, Vincent and Noris 1989, Cilliers 1991, Sharma 1994, thin layer of gold in a sputter coater (Ion-sputter JFC-1100) Sidhu et al. 1994, Hodalova and Martonfi 1995, Nakajima and viewed under SEM. Characters observed included pollen and Monteiro 1995, Kaya et al. 1996, Dawar 1998, Dawar shape, length and width, exine thickness at poles and equator, et al. 2002, Osman 2006, Punt et al. 2007, Zafar et al. 2007, and exine surface features. The measurements were based on Wang et al., 2009). Skvarla and Tuner (1966) studied 15-20 readings from each specimen. The following pollen morphology in different tribes of Asteraceae measurements were taken: (1) Polar axis [P]; (2) Equatorial including Inuleae. Lenis (1971) studied sporoderm pattern diameter [E]; (3) P/E ratio; (4) Colpus length; (5) Pore of some members of subtribe Inulinae (Tribe Inuleae). diameter; (6) Spine length; (7) Interspinal distance; (8) Sharma (1994) in his palynotaxonomical approach of Apocolpium; (9) Mesocolpium diameter and (10) Polar area alpine plants of Western Himalaya reported pollen index [Mesocolpium/ Equatorial diameter]. For each characters of Inula. Dawar et al. (2002) studied pollen character measured, the average measurement of 100 pollen morphology of 22 taxa belonging to Inula (s.s.) and its grains was determined. The terminology used here is that of allied genera from Pakistan and Kashmir. Osman (2006) Erdtman (1952, 1969), Faegri & Iversen (1964) and Walker added pollen morphological data of the tribe Inuleae to the & Doyle (1975) and the acetolysis method is of Erdtman flora of Egypt. (1960). July 2016] POLLEN MORPHOLOGY IN INULA 203 RESULTS AND DISCUSSION KEY TO THE POLLEN TYPE Pollen grains are usually radially symmetrical, isopolar, 1a. Spine with rounded-blunt and few acute apices …… prolate-spheroidal rarely oblate-speroidal (Inula obtusifolia), ...................................................Inula rhizocephala-type mostly tricolporate rarely tetracolporate (Iphiona 1b. Spine with acute-acuminate apices.................................2 grantioides), zonoaperturate and colporate. In all the presently 2a. Tectum densely punctuate or punctuate-fine scabrate in studied taxa, the tectum is echinate and spines are mostly between spines…..............................Duhaldea cappa-type acute-acuminate or sometimes rounded-blunt along with few 2b. Tectum sparsely punctate or psilate-punctuate in between acute apices (Inula rhizocephala). Tectum is either densely spines…………........................... Inula acuminata- type punctuate, scarcely punctate or psilate-punctuate (Inula acuminata) (Figs. 1-3; Tables 1, 2). Amongst all the 3 pollen types, Inula acuminata-type is Based on pollen morphological characters (spine apices the largest one as it has maximum number (nine) of studied and polar length) of Inula and related genera, we have taxa, whereas I. rhizocephala-type is the smallest with I. recognized three major pollen types: (1) Inula acuminata- rhizocephala as the sole representative. This type is type (2) Inula rhizocephala-type and (3) Duhaldea cappa- characterized by the spines with rounded-blunt apices along type. with few acute ones. The remaining two pollen-type i.e., Table 1— Pollen characters of Inula and related genera Name of taxa Polar length [P] (µm) Equatorial diameter P/E* Shape* Aperture Spine apex Spine [E] (µm) base Inula acuminata 29.5(31.3)33 ±0.322 27.5(29.2)30 ±0.214 1.07 Prolate-spheroidal 3-colporate Acute-acuminate Psilate- punctate I. britannica 32.5(34.5)37.5 ±0.591 27.5(31.6)32.5 ±0.794 1.09 Prolate-spheroidal 3-colporate Acute-acuminate Punctate I. falconeri 28.6(31.1)35±0.671 28.6(29)30 ±0.147 1.07 Prolate- spheroidal 3-colporate Acute-acuminate Punctate I. kalapani 25(26.3)28±0.380 25(25.2)25.5 ±0.059 1.04 Prolate-spheroidal 3-colporate Acute-acuminate Punctate I. macrosperma 35(36.2)37.5 ±0.319 32.5(33.6)35 ±0.275 1.08 Prolate-spheroidal 3-colporate Acute-acuminate Punctate I. obtusifolia 31.3(33.8)37.5 ±0.706 35(36.4)37.5 ±0.327 0.93 Oblate-spheroidal 3-colporate Acute-acuminate Punctate I. orientalis 34(34.8)37.5 ±0.574 32.5(33.6)35 ±0.305 1.03 Prolate-spheroidal 3-colporate Acute-acuminate Punctate I. racemosa 35(38.6)41.3 ±0.627 36.4(38.5)40 ±0.344 1.0 Spheroidal 3-colporate Acute-acuminate Punctate I. rhizocephala 35(36.5)40 ±0.584 35(36.4)37.5 ±0.276 1.0 Spheroidal 3-colporate Rounded- Subpsilate blunt+ acute -punctate I. royleana 40(42.7)45 ±0.506 37.5(38.1)38.8 ±0.165 1.12 Prolate-spheroidal 3-colporate Acute-acuminate Punctate Duhaldea cappa 30(31.8)35 ±0.511 27.5(28.9)31.3 ±0.482 1.10 Prolate-spheroidal 3-colporate Acute-acuminate Punctate D. cuspidata 30(32.3)37.5 ±0.750 28.8(30.2)32.5 ±0.387 1.07 Prolate-spheroidal 3-colporate Acute-acuminate Punctate D. eupatorioides 32.5(33.7)35 ±0.298 30(31)32.5 ±0.295 1.08 Prolate-spheroidal 3-colporate Acute-acuminate Punctate Dittrichia 25(25.7)27.5 ±1.125 22.5(25)28 ±0.609 1.02 Prolate-spheroidal 3-colporate Acute-acuminate Punctate graveolens Iphiona 37.5(38.6)40 ±0.294 35(36.3)37.5 ±0.306 1.06 Prolate-spheroidal 3-4-colporate Acute-acuminate Punctate grantioides *P/E=1.00-1.14 (Prolate-spheroidal); P/E=1.00 (Spheroidal) and P/E=0.88-1.00 (Oblate-spheroidal) Table 2—Pollen characters of Inula and related genera. Name of taxa Distance between the apices Equatorial diameter (D) [µm] Apocolpium index [d/D] of two ectocolpi (d) [µm] Inula acuminata 14.5(18.2)22.5±0.233 27.5(29.3)30±0.214 0.62 I. britannica 16(20.9)24±0.081 27.5(31.6)32.5±0.794 0.66 I. falconeri 14(18.9)19.5±0.336 28.8(29)30±0.147 0.65 I. kalapani 11(12.6)14±0.399 25(25.2)25.5±0.059 0.50 I. macrosperma 18(23.5)24±0.278 32.5(33.6)35±0.275 0.70 I. obtusifolia 22(25.5)27±0.714 35(36.4)37.5±0.327 0.70 I.