This article was downloaded by: [University of Bath]On: 09 November 2014, At: 12:01Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH,UK

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Spore Morphology of IndianFerns I. AspidiaceaeB. K. Nayar a & Santha Devi aa Pteridology Laboratory , National BotanicGardens , LucknowPublished online: 01 Sep 2009.

To cite this article: B. K. Nayar & Santha Devi (1964) Spore Morphologyof Indian Ferns I. Aspidiaceae, Grana Palynologica, 5:1, 80-120, DOI:10.1080/00173136409429131

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GRASA PALSXOLOGICA 5 : 1, 1964

SPORE i\IORPT-IOLOGY OF INDTAK FERNS I. ASPIDTACEAE

B1'

U. I<. NAYAR antl SANTHA DEVI

I. JIaterial and JIctliotls . . . . . . . . . . . . . . . . SO 11. Descriptions . . . . . . . . . . . . . . . . . . . . 81

111. Discussion. . . . . . . . . . . . . . . . . . . . . 104 11'. Summary . . . . . . . . . . . . . . . . . . . . . 119

Ilefercnces . . . . . . . . . . . . . . . . . . . . . 119

I. Material and RIetliods

The present study was underta1;en with a view to assess the relia- bility of spore morphoIogy in specific iclcntification of ferns, and to evaluate its significance in comparative studies as a possiblc indicator of the trends of evolution within genera ant1 higher t a n . A preliminary report on the spore morphology of some Japanese Aspidiaceac has been published earlier (Sayar & Dcvi 1 OG3). The present communication ticals with 102 species of tlic Aspitliaccac sensii Copeland 1947 from various pa rk of India.

?'lie inaterial was collected in the ficltl from fresh plants. Specimens of the ferns from which spores werc collcctetl h a w heen deposited in the herbarium of the Xntional IZotanic Gardens, Luckno\\-. Our observations arc haset1 on acetolysed spores mounted in glycerine jelly (Erdtman 1932). For p~irposes of comparison antl in order to facilitate reference by tasonomists, untreated spores niountcd in glycerine jelly were also studied. For each species several spore samples hare been studied. For the sake of lmxity, howcvcr, only one sample is cited for each species.

l'hc genera are arranged according to the classification of Cope-

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S P O R E JIORPIIOLOGY O F I S D I A S FEItSS 81

land (1947). The species of each genus are arranged in alphabetical order. The first species of each genus is described in detail. With regard to the succeeding species only such details in which they differ from the first one are mentioned. In descriptions, details re- garding the material used are given within hrackets against the name of the species in the follo\ving order: reference number of slide, locality, name of collector, and collection number. JIeasurements are averages of a minimum of 10 readings in each plane for each sample, and are expressed as polar axis x longest equatorial dia- meter X shortest equatorial diameter (P X El X E J . Also in the case of concavo-convex spores, the actual length of the polar axis (Fig. 1 a-b) from pole to pole is given as P. In order to give a n idea of the extent of the concavity this reading, as suggested by Ertltman (194G), is followed within bracltets by the distance from the distal pole to an imaginary line connecting the curved ends of the spore on the proximal side (Fig. 1, a-c) minus the distance between this line and the prosinial pole (Fig. 1, b-c). All measurements are exclusive of the perine and any processes of the esinc.

Details regarding the morphology of the spores are quoted in the following order: aperture, symmetry, shape (in lateral and polar view respectively), size, characteristics of laesura, esine, and perine (if present). l’he letters ‘lat’ and ‘pol’ in the descriptions denote the shape of the spore in lateral and polar vie\\. respectively. With regard to the shape in polar view, the terms ‘orate’ (with one end more narrow than the other), ‘oblong’ (ends equal, long sides almost parallel), and ‘elliptic’ (ends tapering equally, long sides conspi- cuously convex) arc used. The colour, measurements, etc. of un- acetolysed spores are recorded within brackets preceded by the letters ‘unacet’. In describing the foldings of the perine, terms such as ‘rugulate’ (with irregular ridges), ‘lobate’ (with thin, ear-like lobes), ‘tuberculate’ (lobcs irregularly circular or oval in outline and having a central depressed area), are used. Tliese terms do not refer to ornamentations of the perine.

The figures of spores in lateral view are reproduced at a magni- fication of 550 times the actual size. those in polar view at 500 times, and sporoderm sections as \veil as ornamentation patterns at 1000 times. The drawings are based on camera lucida slxtchcs. Though eflorts have been made to select a spore as near the average size as possible, the illustrations may not necessarily depict the average size mentioned in the test. 6-631134 G r a m Pal. rol. 5 : 1

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82

11. Descriptions

1. I\ lhTTEUCCIA Totlaro

1. 111. orienfcclis (Hook.) Trer. (8’74; Shillong Peak, Assam; Sayar i5358). Fig. 2 (X.B. the illustrations, 14G figs. in 11 plates, arc reproduced in pp. 103-115).

Spores monoletc, hilateral, plano- to slightly concavo-conves (lat), ovate (pol), 62 x i 0 x 51 p (unncet. 48 X GG X 52 p). Laesura 42 p, tenuimarginate.

Esine 2.2 p thick, light brown. Sesine nearly as thick as ncsinc, smooth. I’crine nearly hyaline, densely granulose, loose, sparsely folded into proniincnt, irregular, thick ridges, most of which arc parallel to the longest equatorial axis of the spore, ant1 protruding about 18 p from the esinc surface. Unacetolysetl spores pale grcenisli- yello\v, with dense granular contents and chloroplastids.

2. DIACALI’E I3lume 2. D. aspidioides I31. (713; Elephant Falls, I<. d J. Hills, Assam; Xayar 7734G). Figs. 3, 105.

Spores monolcte, l~ilnteral, plano- to concaro conws (Iat), elliptic (pol), 32 x 4G X 31 p (unacct. 22 X 34 X 26 p). Laesura 23 p, tcnuimarginntc.

Exine 2.2 p thick, brown. Scsinc nearly as thick as ncsine, smooth. I’erinc light hrown, faintly antl sparsely granulose, loose, folded into thin, convoluted, often irregularly circular or o\-al, ridges, more or less crowded, and protruding about 0 p from the csinc surface. ?‘he folds on the proximal face are often parallel to the lacsura.

3. I’ERXNESIA Don 3. .P. cynfheoidcs Don (525; Elephant Falls, K d J . Hills, Assam; Xayar 57340). Figs. 4, 104.

Spores monoletc, bilateral, plano- to concaro-convex (lat), orate (pol), 23 X 3G X 2G p (unacct. 2G X 34 X 25 p). Laesura 30 p , tenui- marginate.

Esinc 1.0 p thick. brown, not clearly subtlividetl into nesinc and sesine, smooth. I’crine deep brown, faintly antl sparsely granulosc, loose, folded into closely placed thin sinuous ridges, sonic of which are convolute or irregularly circular or oral, protruding about 8 p from the csine surface. Unacctolysed spores pale grecnish-\.cllow, with dense chloroplastids antl with more evident perine granulation.

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SPORE MORPIIOLOGT OF I S D I A S FEIlSS 83

1. ACROPHORUS I’rcsI

4. A. dipellatiis (IVall.) Moore (537; Shillong Peak, Assam; Sagar 64505). Figs. 5 , G, 107.

Spores monolete, hilateral, plano- to coiicaro conws (lat), oblong (pol), 2G X 40 X 26 p (unacet. 20 X 34 X 20 p). Laesura 22 ,LC,

tenuimarginate. Esine 1.G p thick, light brown, not clearly subdiridctl into scsine

and ncsine, smooth. Perinc pale Inown (unacct. darker), protruding about G ,u from the esine surface, densely granulose, loose, sparsely folded into thin irregular ridges, most of which are parallel to the equatorial asis of the spore. Unacctolysed spores with dense, pale yellow plastids.

5. HYPO D E 1 I A T I U 11 Kunze

5. H. crencifrcni (Forsk.) Iiuhn (10-16; Panchmarhi; J. G. Srirastara 58519). Fig. 7.

Spores monolete, bilateral, plano- to concaro-convcs (lat), ohlong (pol), 32 X 45 X 32 p (unacct. same). Laesura 28 p, crassiniarginalc (with smooth thin I ips).

JSsine 2.2 p thick, reddish brown (unacct. same). Scsinc nearly as thick as ncsinc, smooth. I’erinc deep brown (unacet. same), protruding ahout G p from csinc surface, densely granulosc, loose, foldctl into thin, short, often sinuous, lobatc, more or less sparse ridges.

G. I’OI,YSTICHU1I Roth.

6. P. ncnnlhophyllrmi (Franch.) Christ (1 174; Tonglu, Darjeeling, Dengal; P. Chnndra G0113). Figs. 10, 11.

Spores monolete, hilateral, plano-conrcs (Iat), oblong (pol), 24 X 3G x 28 p (unacet. sanic). Laesura 20 p, tenuimarginate.

Esim 1.5 p thick, light brown (unacct. same), not clearly sub- tliritletl into sesinc arid nesine, smooth. I’erine deep bro\vn (unacet. darker), protruding up to 4 p from the csinc surface, more or less loose, minutely spinulosc, highly wrinkled into subconical lobes \vhich are densely aggregated together. Lobes with more or less pointed apices, about G p (4-8 p) across at the base. Uiiacctolysctl spores with sparse pale green chloroplastids.

7. P. acidentum (L.) Roth. (569; Snodon, I<. c!! J. Hills, Assam; Xayar ’77306). Figs. 15-17.

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84 I%. I<. SAP.\R A S D SASTIIA DEVI

Spores 37 X 52 x 3G p (unacet. same). Laesura 44 p. Esinc 3.3 p thick. Scsinc nearly as thick as ncsinc. Perine protruding up to 13 p from esinc surface, granulose; lobes subconical, blunt, about S ,LL (4-12 p) across at base.

8. P. acitleaticni var. nirccronifoliicni U1. (1 172; Sikkim; P. Chantlra 97802).

Spores 32 X 46 X 30 p (unacet. same). Laesura 22 p. Esinc 2.6 p thick, pale brown. Sesine nearly as thick as nesine. Perinc dark brown (unacct. light brown), prominently granulose; folds protruding up to 12 p from esine surface, hemispherical to subconical, about 10 p (8-12 p) across at base.

9. P. aiiriciilutiini (L.) l’r. (247; Chcrrapunjee, Assam; Kayar 49Si-1). Fig. 110.

Spores plano- to & concavo-convcs (lat), 28 x 41 x 30 p (unacet. 24 x 36 X 24 p). Laesura 24 p. Esirie 2.2 p thick. light brown. Perine brownish, densely granulosc, protruding up to 5 p from mine surface; folds with thin pointed apes and more or less bulbous base, G p (4-8 p) across at hase.

10. P. biaristatmn Moore (1044; Pynursla, I<. S: J. Hills, Assam; Xayar 77331). Fig. 108.

Spores plano- to concaVo-conws (lat), elliptic (pol), 34 x 48 x 37 p (unacet same). Laesura 33 p. Esinc 2.5 p thick. Scsine nearly as thick as ncsinc. I’crine light brown, faintly granulose. Folds of perinc protruding up to 12 p from esinc surface, subhemispherical to bluntly subconical, 8 p (4-16 p) across at the base.

11. P. cfiscrcfizm (Don) Dicls (1173; Tonglu, Darjeeling, Bengal; P. Chandra 69405).

Spores 21 X 32 x 26 p (unacct. 20 X 26 x 20 p). Laesura 20 p. Esine 1.0 p thick. Perine pale hrown (unacct. same); folds about 2 p across at the haw, generally crowned hy one or two spinulcs, and protruding up to 4 p from esine surface.

12. P. lerititni (Don) XIoore (1019; Cherrapunjee, I<. & J . Hills, Assam; Kayar 50465).

Spores 20 X 30 x 22 p. Laesura 16 p. Esinc 2.2 p thick. Sesine nearly as thick as nesinc. Perine hrown, densely granulosc and sparsely spinulose; foldings protruding up to G p from the mine surface, nearly liemispherical, about 6 p (4-S p) across at the hasc.

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SPORE JIORPIIOLOGT O F I S I I I A S F E R S S 85

13. P. lobatrim (Huds.) Pr. (SGS; llahabaleshwar, lfaharaslitra; R. X. Ghosli 97SO-I). Fig. 14.

Spores ovate (pol), 26 x 39 x 25 p. Laesura 26 p. Esine 2.2 p thick, golden brown. Sesine nearly as thick as ncsinc. Perine golden hrown, loose, faintly granulosc. sparsely folded into large, irregular, lobate ridges, most of which are parallel to the equatorial asis of the spore, and protruding up to 8 p from the esine surface.

14. I] . loncliifis (L.) I\otli (1148; l‘hatwas glacier, liaslimir; G. S. Srivastava l l G 7 7 ) .

Spores 24 X 36 X 28 p (unacct. same). Laesura 1 G p. Esinc 1.5 ti thick, deep lirown (unacct. same). Scsinc nearly as thick as nesine. Perine adherent to csine, spinulose; folds subconical, protruding up to 4 p from tlic isine surface, usually c r o w m i liy a sharp spinule, about 3 p across at the base, appearing like spines with bulbous bases.

15. P. ntarginafttm Wall. (1171; Bihar; J. G. Srivastara 60610). Spores ovate (pol), 36 x 46 x 30 p (unacct. same). Lacsura 24 p.

Esine 2.5 p thick. Sesine nearly as thick as ncsine. I’crinc granulose; folds nearly hemispherical, 8 p (4-12 p) across at base, and protrud- ing up to 8 p from the esine surface.

16. P. obliqiziznz (Don) ;\loore (1 112; Iiaslimir; G. S. Srivastara

Spores 32 X 48 X 32 ti (unacct. same). Lacsura 24 p. Esine 2.2 p thick, deep brown (unacet. same). Sesinc nearly as thick as nesine. Pcrine granulosc; folds hemispherical to subconical, about 20 p (8-28 p) across at the base, and protruding up to 16 11 from esine surface.

10 188).

1 7 . P. speciosrim (Don) J. Sm. (77; Bombay Shola, Kodail;anal, Madras; Kayar 44485). Fig. 109.

Spores orate (pol), 33 x 46 x 37 p. Laesura 24 p. Esine 1.6 p thick. Sesine nearly as thick as ncsinc, deep brown. I’erinc l~ro\vnish, more or less adherent to the csine, granulosc; folds small, conical, up to 4 p across at base, with pointed apex, and protruding up to 4 p from esinc surface.

18. P. sqirarrosum (Don) F6e (100; I’ycara. Ootacaniund, XIadras; Xayar 45524). Figs. 12, 13, 137.

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86 l3. I i . SAYAR A S D SSSTIIA DEVI

Spores plano- to concavo conves (lat), elliptic (pol), 34 x 50 x 37 p (unacct. 27 x 40 x 28 p). Laesura faintly crassirnarginate, 43 p. Esinc 2.8 p thick. Sesinc thiclicr than nesine, pale yellowish hewn (unacet. same). Perinc pale yellowish brown (unacet. same), tlcnscly granulose, folded into irregular thin lobate ridges, ant1 protruding about 10 p from esine surface.

19. P. stinirilans Pr. (1110; Tiksain, Kashmir; S. Singh. 53334).

Spores 24 x 3,3 x 27 ,u (unacet. same). Laesura lop, crassimarginatc. Esinc 2.2 ,u thicli, pale brown (unacct. lighter). Sesinc nearly as thick as nesinc. I’crinc pale brown (unacet. lighter), densely granulose, morc or less sparsely folded into elongated large lobate ridges, and protrntling up to 16 ,u from the esine surface. I’crine foldings less prominen2 (nearly absent in some) in unacetolyscd spores.

7. LITHOSTEGIA Ching

20. L. foeniculacca (Hooli.) Cliing (508; Pynursla, I(. L‘ J. Hills, Assam; Sayar 75314). Figs. 8, 9, 10G.

Spores monolcte, bilateral, plano- to concaro-convex (lat), oblong (pol), 28 x 44 x 29 p (unacet. 24 x 36 x 24 p). Laesura tenuimargi- nate, 2s p.

Esine 1.0 p , thick, (lark-brown with a rcddish tinge, not clearly clifTercntiated into scsine and ncsine, smooth. Perine dark brown (unacct. darker), faintly granulose (unacet.: granulation not evident), loose, protruding up to 8 p from mine surface, and folded into a few, elongated, thick, conrolutcd ridges which may be irregularly circular or oral and rarely crowded.

Fig. 18.

8. CTRTO;\IIU;\I I’resl

21. C. cargotidcorn (Wall.) I’r. (635; Bagutliar, Almorah, Uttar I’radesh; J. G. Srivastava 53720).

Spores monolcte, bilateral, plano- to concaro-conres (Iat), oblong (pol), 37 x 51 x 12 ,u (unacet. same). Laesura 12 p, tenui- marginate.

Esixic 1.5 ,u thick, light brown (unacct. same). Scsinc nearly as thick as ncsine, smooth. Perine brown (unacet. same), smooth, densely folded in a tuberculate pattern, and protruding up to 4 p from esine surface.

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Sl’OI\E SIORPIIOLOGT O F I S D I A S FEHSS ST

22. C. fdccrti im (L. fil.) Copel. (1170; Cult. X.U.G.). Fig. 21. Spores 32 x 45 x 32 p (unacct. same). I’erine h o w n (unacet.

darker). Foltlings of perine less prominent, smaller tlian in C. cargo- fi t leuin, anti protruding up to 4 p from esinc surface. Unacctolysed spores with pale chloroplastitls.

23. C. hookerianum (I’r.) C. Chr. (1045; Cherrapunjcc, Assam; Nayar 7i34S). Figs. 19, 20, 111.

Spores 24 X 40 X 32 p (unacet. same). Laesura 28 p. Esine 1.3 p thick, not clearly differentiated into sesine and nesine. I’erine dark hron-n (unacct. same), densely granulosc, densely foltlcd into hemi- spherical to hluntly conical folds which are G p (4-S p) across at base, protruding up to S p from esinc surface, and aggregated so as to form a pattern similar to that in Polysticliiiiii spp.

9. RUlIOHI\A Raddi 21. R. crrislafa (Forst.) Ching (862; Agumbe, Shimoga, llysore; Xayar 45551).

Spores monolete, bilateral, plano-convex (lat), oratc (pol), 30 x 42 X 26 p (unacet. 26 x 34 x 26 p). Laesura 17 p, tcnuimar- ginate.

Esine l.G p thick, light hewn (unacet. same). Sesinc nearly as thick as nesinc, smooth. Perine light brown (unacet. same), faintly granulose, closely adherent to esine, sparsely folded into thin, elongated, irregular ridges, and protruding up to 1G p from the cxine surface.

25. R. carvifolicr (Icze.) Cliing (289; Shillong I’cnk, Assam; Nayar cC: Prcm La1 50413). Figs. 22, 1 1 1 a.

Spores elliptic (pol), 24 x 28 x 21 p (unacet. same). Laesura 24 p. Esinc 2.2 p thick. Sesine thinner than nesine. Perine nearly liyalinc (unacet. same), prominently granulose, loose, sparsely folded into elongated thin ridges protruding about 10 p from the esinc surface and mostly parallel to the equatorial asis of the spore.

10. I30LI3ITIS SCllOtt 2G. B. costafa (Wall.) Ching (551; l‘ura, Garo Hills, Assam; Xayar 50032). Figs. 25, 112, 113.

Spores monolcte, bilateral, plnno-conrcs (Iat), oratc (pol), 32 x 45 x 35 p (unacct. 24 x 3G X 2s p). Laesura 29 p , tcnuimar- gi na t e .

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Exilic 1.0 p thick, liglit b r o \ ~ n (iinacct. deep brown), not clearly cliffercntiatctl into scsine antl ncsine, smooth. I’crine light hrown (unacet. deep l)ro\~n), densely but faintly grnnulose, loose, folded into thick, irregular, short ridges sonic of which forni bluntly conical protuberances, and prolrucling up to 8 p from esinc surface. The lacsura in sonic spores is forltctl at one end and rarely appears trichotomous (Fig. 1 13). Unacetolysed spores with dense, globose, pale green chloroplasts.

27. B. crispcrfulrt (V‘all.) Cliing (52G; ‘rum, Garo Hills, Assani; Xayar 63414). Figs, 29, 114.

Spores 30 x 38 X 32 p (unacct. 2s X 32 X 28 p). Laesiira 28 p. I’crinc very sparsely folded into faint folds (sometimes folds nearly absent) most of which are paralIe1 to tlie equalorial axis of the spore and protruding up to 12 p from tlic csinc surface.

28. B. tliuersifolici (Fr.) Cliing (590; Cult., Singapore Isot. Gard.). Spores 32 x 48 X 38 p. Laesura 26 p. Esinc 3.0 p thick. Sesinc

nearly as thick as nexine. I’crinc brownish with a 1)lacltisIi Iiuc, prominently graniilose, highly foltlccl into elongated, thick, more or less sinuous ridges protruding up to 19 p from the esine surface.

29. B. heferoclitcr (Pr.) Ching (51 8; Darugiri, Garo Hills, Assam; Nayar G3440). Fig. 31.

Spores oblong (pol), 28 X 38 X 30 p (unacet. 21 x 32 2G p). Laesura 22 p. Thine 3.3 p thick, brownish (unacct. same). Sesinc ncarly as thick as nesine. I’erine brownish (unacet. same), smooth, highly foltlctl into thin, more or less sinuous, elongated ridges most of which radiate from the centre to the periphery (in lateral Tien- of the spore) antl protrude up to 12 p from the mine surface.

30. B. prcslicinci (Fee) Ching (iG8; Agumbc, Shimoga, Jlgsore; 1’. Chandra 77712). Figs. 23, 24, 116.

Spores 28 X 37 X 30 p (unacct. 2G x 32 x 28 p). Lacsura 24 p. Esinc 1 .G p thick. I’erine bro\~nish (unacet. yellowish brown), irregularly rugulate, more or less atlliercnt to the esine and scarcely folded.

31. 13. seniicordcrfa (Aloorc) Cliing (792; Jog Falls, Nysorc; 1’. Chandra 75727). Figs. 26-28, 120.

Spores oblong (pol), 30 x 40 X 3 G p (unacet. same). Lncsurn 22 p. Esine 1.6 p thick. Perine dark brown (unacet. same), prominently

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SPORE ~IORI’ I IOLOGS OF I S D I A S 1:EIiSS 89

granulose, more or less adhcrcnt to the esine, folded into small, nearly semicircular or subconical folcls (aggregated together and inore or less regularly arranged) and protruding up to 8 p from the csine surface.

32. 11. wibcrenata (Hook. ct Grey.) Ching (97; I<anchciikumri, Sakleshpur, JIysore; Sayar 45559). Figs. 32, 11 7.

Spores 34 x 37 X 32 p (unacct. 28 X 34 X 30 p). Laesura 31 p. Esinc 2.2 p thick. Sesinc nearly as thick as ncsinc. I’erine highly wrinlilcd into thin; sinuous, elongated ridges, some of which appear lobatc, and protruding up to 10 p from the esinc surface.

33. B. subsiniplex (Fk) Ching (407; Tura, Garo Hills, Assani; Sayar 6 I’rem La1 50931). Figs. 33, 115.

Spores 23 x 34 x 27 p (unacet. same). 1,aesura 24 p. Exilic 1.6 ,u thick, tlarli brown (unacet. darker). Perine brownish with a l h c k hue (unacet. darker), highly wrinlilecl into thin elongated ridges, protruding about 12 ,u froni esine surface and mostly radiating from centre to periphery in the lateral view of spore. Itarely the ridges may anastornose forming irregular reticulations. Prominent plastids and few ycllowish oil globules occur in unacetolysed spores.

34. B. uircns (Wall.) Schott (479; I’onmudy, Trivandrum, Kernla; Sayar 45058). Fig. 30.

Spores 30 X 43 X 38 p (iinacet. 26 X 35 X 28 p). Lacsura 21 p. Esine 1 .G p thick, pale brown (unacct. yellowish brown). Pcrine light brown (unacet. same), highly folded into thin, nearly semi- circular, aggregated ridges, some of the nearby. ones of \vliicll coa~escc to form elongatetl unciulating ridges (unacet.: ridges sliarpcr and more irregular), and protruding up to 8 p from the csinc surface. Unacctolyscd spores with prominent plastids and i~sually one large central golden-yellow oil globule.

11. EGESOLFIA Scliott

35. E. crppendiculata (\\‘ilIcI.) J. Sm. (1 72; ;\Iercara, Coorg, JIysorc; Xayar 45555). Figs. 37, 121.

Spores monolcte, I>ilateral, plano-conrcs (Iat), elliptic (pol), 34 X 45 X 36 ,u (unacet. same). Lacsura 34 p , tcnuimarginate.

Esine 1.6 p thick, light brown (unacet. same). Sesine nearly as thick as nesine, smooth. Perine Inown (unacct. same), prominently

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reticulate (reticulation fine), loose, lightly folded into many elongated sharp ridges, most of which radiate from the centre of the spore (in lateral +w), and protruding up to 20 ,LJ froin the esinc surface. Unacetolysetl spores contain dense pale green chloroplastitls.

36. E. asplenifolirt (Bory) Fde (107; Ponmudi, Trirandriim, Kernla; Xayar 45057). Figs. 38, 39.

Spores oblong (pol), 31 x 49 x 37 p (unacet. 28 x 36 x 28 p). 1,aesura 32 p. Esinc 2.8 p thick. Scsinc thicker than nesine. I’crinc hen-n (unacct. dark brown), protruding about 12 p from csinc surface, distinctly reticulate (muri thickened at the corners).

37. E. helferiancr (I<ze.) C. Chr. (742; Parlobjig, JIiddle Xndamans; G. S. Srivastara 79381).

Spores plano-conres to more or less biconres (lat), 3G X 4i X 36 p. Lacsura 37 p, its margins faintly incrassate. Esine 2.2 p thick, dark brown. Scsinc conspicuously thiclter than nesinc. Perinc dark 1)row-n; its ridges more or less angular, protruding up to 11 p from mine surfacc.

35. E. sinensis (I3ak.) ,\lason (543; Darugiri, Garo Hills, Assam; Xayar 63431). Figs. 3-1-36, 119.

Spores ovate (pol), 32 x 35 x 32 p (unacct. same). Iaesura 24 p. Esine 1.0 p thick, 1xown (unacet. dark brown). I’erinc light brown (unacet. darker), its reticulation rather obscure. One or two ridges of pcrine arc elongated parallel to the longest equatorial asis.

39. E. uiuipara (Hook.) C. Chr. (560; Tura, Garo Hills, Assam; Xayar 50028). Figs. 40, 118.

Spores plano-conres to more or less hiconves (lat), oblong to short and nearly circular (pol), 32 x 42 x 35 p (unacct. 26 x 32 x 28 p). Laesura 20 p. Esinc about 2.2 p thick. Pcrinc \rrinltled into thin angular ridges protruding up to 20 p froin esinc surface. Some ridges coalesce, forming irregular reticulate patterns.

12. ELAPHOGLOSSUAI Schott

40. E. conforme (Sw.) Schott (531; Shillong Peak, Assam; Xayar (34508). Figs. 41, 122.

Spores monoletc, bilateral, concaro-convcs (lat), clongate-oblong (pol), 28 (36-8) X 57 x 33 p (unacet. 23 X 40 x 28 p). Laesura 21 ,LL, margins faintly incrassate.

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SPORE MORPIIOI.OGP OF I S D I A S FERSS 91

Esiiic 1 .G p thick, light brown (unacet. same). &sine tliiclxr than ncsinc, smooth. I’crinc hewn (unacet. darker), distinctly granulose (some of the granules coalcscc to form riigulac or incomplete reti- culations), loose, wrinkled forming more or less sparse, small, conical projections or short, undulating ridges, protruding up to 12 , ~ i from the csinc surface.

13. DRYOPTERIS Xdanson

41. I). burbigem (JIoore) 0. I<tze. (1116; Elephant Falls. I<. B J. Hills, Assam; Xayar 40SS3). Fig. 43.

Spores monolete, bilateral, plano-conrcs to inore or less concavo- conres (Iat), elongate-oblong (pol), 30 x 48 x 30 p (unacet. same). Lacsura 24 p, tenuimarginate.

Esinc 1 . G p thick, light brown (unacet. same). Scsine nearly as thick as nesine, smooth. Perine brown (unacet. same), very faintly granulose (unacet.: granulations not apparent), \rrinklcd into short, irregular, often curved, lobate, sometimes tubcrculatc, crowded ridges, and protruding up to 8 ti from esine surface.

42. I). coclilecifa (Don) C. Chr. (i32; Parasnath, Ililiar; 1‘. Chandra 34074). Fig. 126.

Spores elliptic (pol), 37 x 54 x 41 p (unacet. 31 x 48 x 28 p). Laesurn 33 p. Esinc deep brown. Scsinc slightly thicker than nesinc. I’criiic wrinkled into thin, irregular, often angularly curved, zigzag folds which are often lobate, and protruding up to G p from the esine surface.

43. D. fi2i.r-mas (L.) Schott (30G; Sliillong Peak, I<. B J. Hills, Assam; Sayar 773Gi). Figs. 42, 123.

Spores oblong (pol), 32 x 49 x 82 p (unacet. 24 x 3G x 28 p). Laesura 32 p. Esinc brown, not distinctly difierentiatcd into sesine and nesine. I’erine deep brown (unacet. darker), smooth, loose, wrinkled into broad, cro\\-ded, zigzag, sometimes angular ridges protruding up to 12 p from the esine surface.

44. D. h i r f i p s (I31.) 0. Ktzc. (575; Shillong Peak, I<. L! J. Hills, Assam; Kayar G4502). Figs. 44, 124.

Spores plano-conres (lat). elliptic (pol), 30 X 47 X 32 p (unacet. 23 X 31 X 24 p). Laesura 36 p. Esine 2.2 p thick. Perine smooth, wrinkled into many irregular, thin, crowded folds some of which may he lohate.

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45. D. sparsa (Don) 0. I<tze. (523; Mudigere, Iiadur, JIysorc; Xayar 45571).

Sporcs plano-convex (lat), 45 X G i X 50 p. Laesura 32 p. Pcrinc (lark brown, smooth, wrinkled into distinct, short, irregular, thin, often angularly folded, crowdcd ridges, and protruding up to G p from the esinc surface.

14. CTESI1‘IS C. Chr.

16. C. licndersoriii (Bctld.) Copel. (5G4; Cult. S.B.G.) . Figs. 45, 129. Spores monolete, bilateral, plano-convex to more or less concaro-

convcs (lat), oblong (pol), 28 x 41 x 32 p (unacct. 22 x 3G x 24 p). Laesura 24 p, tenuimarginate.

Esinc l.G p thick, deep hewn (unacct. same). Sesine thicker than nesine, smooth. I’erinc deep brown (unacct. same), smooth, more or less loosc, folded into thin, sharp, very sinuous ridges, sonic of which appear lobatc or even tubcrculate (unacet.: folds more promi- nent), and protruding about 1 p from the csine surface.

15. TECTXRIA Car.

17. T. cicrrtaricr (L.) Copd. (730; Girnar, Sanrashtra; Janki Prasad 14054). Figs. 4G, 135.

Spores monolctc, bilateral, plano- to faintly concaro-conrcs (Iat), oblong (pol), 32 X 44 x 32 p (unacct. 24 x 3G x 24 p). Lacsura 24 p, tenuimarginate.

Exinc l.G p thick, light brown (unacet. same). Sesine nearly as thick as nesine, densely granulose. I’erinc nearly hyaline (unacet. same), dcnscly gmnulosc, loosc, sparsely folded into a few zigzag, elongated folds (most of which are parallel to the longcst equatorial axis of the spore) and protruding u p to 8 p from the mine surface.

45. T. deuexa (Iize.) Copel. (358; Chcrrapunjee, Assam; Kaym 50912). Figs. 49, 50, 130.

Spores plano-convex (lat), 24 x 37 X 27 p (unacet. same). Laesura 28 p, its margins faintly incrassate. Esinc 1 p thick, not distinctly subdirided into scsinc and nesine, smooth. I’erinc light brown (unacet.: ncarIy hyaline), densely spinulosc, sharply foltleti into more or less sparse, irregularly distributed lobes (some of which are angularly folded), and protruding u p to G p from the esine surface.

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SPOllE JIORPIIOLOGY OF I S D I . \ S F E R S S 03

49. T. fitscipes (U‘all.) C. Chr. (850; Tura Ridge, Garo Hills, Assarn; Xayar 50999). Figs. 47, 48, 133.

Spores plano-convcs (lat), 30 x -13 x 28 p (unacet. 28 x 40 x 26 i t ) . Laesura 28 p. Esine 2.2 p thick, brown (unacet. yellowish hrown). Sesine thicker than nesinc. Pcrine nearly hyaline (unacct. pale- yellowish), spinulose, sparsely folded into thin, irregularly distri- buted, sinuous ridges, and protruding up to 8 p from tlic esinc surface.

50. T. gigantea (Bl.) Copel. (115.i; I’alghat, Jlalabar, Kerala; Xayar 455G1).

Spores plano-conves (lat), ovate (pol), 2G x 41 x 28 p (unacet. same). Laesura 20 p. Esine 2 p thick, not clearly subdivided into sesinc arid nesine. I’erinc wry light hroivn (unacet. darker), closely adhering to the esine, spinulose, wrinkled into irregular, sinuous thin ridges, most of which run in the samc direction as the polar axis of the spore and protrude up to 8 p from esinc surface. Spinulcs thin, sharp, nearly hyalinc.

51. T. heferosorci (Baker) Ching (1 168; I’unalur, Kcrala; Nayar

Spores plano-conves (lat), 32 x 43 X 31 p (unacet. 26 x 38 x 26 p). Lacsnra 23 p. Esinc 2.3 p thick. Scsinc slightly thicker than nesine, faintly granulose. I’erinc pale brown (unacet. darker), adherent to the esinc, densely spinosc (spines thin, slender, pointed, often curved, 8 [L long), folded into thin; faint ridges connecting the bases of the spines and thus forming a n irregular reticulate pattern.

52. T. macrodonfa (Fk) C. Chr. (552; Snodon, I<. 6 J. HilIs, Assam; Xayar 63500). Figs. 54, 55.

Sporcs plano-conves (lat), 32 x 46 x 32 p (unacet. same). Esine 2.2 p thick, deep brown (unacct. darker), densely spinulose. Perine sparsely spinulose, folded into elongated, thin, sparse, irregular, often curved ridges, and protruding about 0 p from the esine surface.

53. T. polyrnorpha (Wall.) Copel. (13; ‘ h r a Ridge, Garo Hills, Assam; Nayar S: I’rem La1 50495). Figs. 51-53, 134.

Spores plano-conves (lat), 28 X 44 X 30 p (unacct. same). Laesura 31 p, its margins faintly thickened. Esinc 2.2 p thick, brown (unacet. darker). Sesine considerably thicker than nesine, densely spinulose. the spinules having sharp apices. Perine smooth, folded into a few elongated thin ridges protruding up to 8 ,u from esine surface.

455G2).

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54. T. semibipiiinrcfn (Wall.) C. Chr. (849; Shengurni I~eserve, I<ulatlioopuzha, Kerala; Xayar 44SlO). Figs. 56, 5 i , 131.

Spores plano-convex (Iat), 27 X 30 X 26 p (unacct. same). Lacsura 20 p. Esine 2.2 p thick, light 1)rown (unacet. same). Sesiric thiclcer than nesinc, smooth. I’erinc spinose (spines uniformly distributed, 4 p long), closely adherent to the esinc and indistinguishable escept at the regions where it forms thin ridges connecting the bases of the spines and thus forming a reticulate pattern over the csine.

55 . T. sinionsii (Baker) Ching (GOO; Turn, Gar0 Hills, Assam; Xayar 63413). Figs. 58, 59.

Spores 28 x 40 x 28 p (unacet. 24 x 32 x 24 p). Laesura 24 p. Sesinc thicker than ncsine, spinulosc. I’crinc sparsely spinulosc (spinulcs smaller than those on the esine), with a few sharp, light folds (which often form angular projections) protruding up to 4 p from tlie esinc surface.

5G. 7’. siibconflitens (Bedd.) Ching (72; Turn, Gar0 Hills, Assam; Xayar 50927). Figs. GO, G1, 132.

Spores plano-convex (lat), 25 x 41 x 31 p (unacct. same). Lacsura 20 p, crassimarginate with thin smooth lips. I<sinc 2.2 p thick, brown (unacet. same). Sesinc thicker than nesine, tlcnsely spinulose. I’erinc hyalinc to pale brown (unacct. same), sparsely and faintly granulosc, folded into large, sparse, lobate ridges protruding up to 0 ,LL from the esine surface.

57 . T. uariolosci (Wall.) C . Chr. (469; 13asistasram, Kamrup, Xssam; Sayar & I’rem La1 50003). Figs. G2, 13G.

Spores elliptic (pol), 30 x 45 x 30 p (unacct. same). Laesura 28 p, Esinc 1 .O p thick, not clearly subdivitletl into scxinc and nesinc, smooth. I’erinc smooth, folded into thick, clongatecl, sinuous, often crowded ridges (some of which are curved, s o n i c h e s tubcrculate) protruding up to 14 p from the esine surface.

5s. T. ruightii (Clarke) Ching ( 1 165; XInliadcr, I<. & J. Hills, Assam; Xayar 77369).

Spores 2G X 38 X 28 ,u (unacet. 21 X 20 X 22 p). Lacs~ira 24 p. Esinc 2 p thick, smooth. Pcrinc spinulosc, wrinkled into short, thin, irregularly disposed ridges protruding up to 8 p from mine surface.

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Sf’OIiE JIORPIIOLOGP OF I S D I I S FEIiSS 05

1G. QUERCIFILIX Copel.

50. 0. zeilcririca (Houtt.) Copel. (630; Cult. Hort. I3ot. Hoenensis). Fig. G3.

Spores monolete, bilateral, plano-conws (lat), elliptic (pol), 3 j x 5 G X 3G 11. Laesura 24 p, tenuimarginate.

Esine 1.G p thick, Inown, not clearly subdivided into sesine and nesine, smooth. Perine more or less loose, deep brown, smooth, wrinkled into irregular ridges (forming a lophate pattern) protruding up to 4 ,LL from esinc surface.

17. L.4S‘IXE.A Uory

G O . 1,. ccrlcaratcc (D1.) Jloore (1087; I‘alparai, Coimbatorc, JIatlras; Xayar 45382). Figs. G4, GS.

Spores monolete, bilateral, plano- to slightly concavo conres (lat), elliptic (pol), 20 x 3G x 24 p. Laesura 16 p, tenuimarginate.

Esine 2.0 p thick, dark brown. Sesine nearly as thick as nesine, smooth. I’erinc dark brown, closely adherent to esine, granulose, sparsely folded into thin, short, irregular ridges protruding about 4 ti from the mine surface. The ridge over the laesura is nearly straight, prominent, and as long as the laesura.

G1. L. cilicrirc (\\‘all.) Hook. ( I 143; Cult. N.13.G.). Spores plano conves (lat), oblong (pol), 30 X 38 X 28 p (unacet.

same). Laesura margins faintly incrassate. Esine brown (unacet. pale brown), not distinctly subdiridcd into sesine and nesine. Perine closely adherent to csine, densely spinose, tleroitl of folds. Spines 4 p long, sharp. Unacetolysecl spores hare dense, nearly colour- less, large plastitls.

G2. 1,. crcrssifo/icc (Bl.) 0. Ktzc. (1144; Cult. X.13.G.). Fig. GG. Spores ovate (pol), 28 x 3G X 25 p (unacet. same). Laesura 12 p.

Esinc 1.G ,u thick. I’erine rery faintly granulose, more or less adherent to esine, wrinltlctl into thin, short, crowded, zigzag, ridges (irregularly disposed escept tlie one over the lacsura which is long and almost straight), protrutliiig up to G p from the esine surface.

G3. 1,. fcrlcilobcc Hook. (G59; Panchmari, Jlarlhya Pradesh; Hira La1 33210). Figs. G i , 65, 125.

Spores plano-conves (lat), oblong (pol), 30 X 40 X 31 p (unacet. same). Laesura 24 p. k i n e 1.G p thick, light hrown, spinose. Spines

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96 11. I<. SAPAR X S D S l S T I I d DEVI

deep brown, 4 p long, thick, crowded, uniformly distributed, tapering from a broad base to a sharp apes, occasionally curved. I’erine absc 111.

61. L. grrccilescens (I31.) Moore (1142, Cult. X.B.G.). Figs. G9, TO. Spores plano-conres (lat), orate (pol), 26 x 38 x 28 p (unacet.

same). Lncsura 18 p. Perinc very closely adherent to esine, smooth, folded into spinose l o h s (some of which may he hluntly conical) protruding u p to 4 ,u from the esine surface. The ridge over the laesura is flap-like, sometimes interrupted. 66. L. ochfhocks (Kze.) Jloorc (1 114; Dcvicolam, Kottayam, Kerala; S a p r 45295).

Spores plano-convex (lat), oblong (pol). 28 X 38 X 28 p. Laesura 20 p. k i n e 2.2 p thick, spinose. Spines hlunt, dense, 4 p long, sometimes group-wise coalescing to form rugulac. Periric absent.

G G . L. phegopferis (L.) Uory (1058; 13clgium; A. Lawalrce 11 900). Fig. 51.

Spores elongatc-ovate (pol), 24 X 38 X 24 p (unacet. same). Laesura 20 p. Esine hrou-n (unacet. nearly hyaline). I’erinc dull brown (unacet. hgalinc), more or less loose, folded into sparse, thin, rugulate ridges. Ridge over the laesiira very distinct, unbroken. Unacetolyscd spores with sparse, pale green plastids of varying sizes and one or two nearly colourless oil globules.

67. I,. piilviniiliferrc Ecdd. (314; Shillong I’cak, I(. CE: J. Hills, Assam; Kayar t? Prcm La1 50437). Figs. i 2 , 127.

Spores oblong (pol), 28 X 36 X 32 ,u (unacet. same). Lacsura 24 p. Esinc l . G p thick, not clearly differentiated into scxine and nesine. Pcrinc smooth, inore or less adherent to the esine, folded into large, thick, lobate, sometimes curred folds protruding up to 4 ,u from the csinc surface.

68. L. pyrrhorhachis (Kzc.) C. Chr. (605; Shillong Peak, I<. 62 J. Hills, Assam; Xayar 64501). Figs. 73, i4, 128.

Spores oblong (pol), 24 X 40 X 26 p (unncet. same). Laesura 28 p, its margins incrassate. Esinc 2.8 p thick, yellowish brown (unacet. nearly hyaline), bearing elongated, sparse, smooth, narrow ridges which often form an irregular large reticulate pattern especially on the distal face. Perine yellowish brown (unacet. nearly hyaline), smooth, closely adherent to the esine and devoid of folds. Un- acetolyscd spores contain prominent plastids of varying sizes.

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SPORE JIOI1I’IIOI.OGT 01‘ I S D I I S FEIlSS 0i

69. L. fenericcridis (Il‘dl.) lloorc (1082; Cult. K.1j.G.). Spores oblong (pol), 24 X 38 X 2G p. 1,aesura 20 p. Esine l . G CL

thick, brownish. Sesine thicker than ncsine. Perine dark hrown, more or less atlliercnt to the csine, prominently granulose (granula thick, wart-like).

15. CYCLOSORUS Link

iO. C. criiiipes (Hook.) Ching (121; Sliangila IZcscrvc, Ihlathoo- puzlia, ICcrala; Xayar 41Gi4).

Spores nionolctc, hilateral, more or less concavo-conses (lat), oblong (pol), 2G (28 - 2) x 4G x 2G p (unacct. 24 x 32 x 2 1 p). Lacsurn 24 p, tenuimarginate.

k i n e 2.0 p thick, brown (unacet. clarltcr). Scsinc tliiclter than riesinc, tlenscly spinulose. Spinules 2 p long, sharp, brown. I’crine absent.

51. C. cristccta (L.) Ching (614; Cult. X.U.G.). Figs. 75, 138. Spores plano-conrcs (lat), 25 X 39 x 31 p (unacct. 24 X 32 X 2-4 p).

Sesine thinner than nesinc. smooth. Perine deep brown (unacet. same), closely adherent to the esine, faintly granulose, lightly folded into thick, short, irregular, often curved ridges (some of which may coalescc and form irregular patterns) protruding up to 4 p from the esinc surface. Ridge over lacsura prominent, elongated.

72. C. didynosora (Parish) C. Chr. (3%; Tura, Garo Hills, .\ssam; Xayar G!! I’rcni La1 50920). Figs. 7 G , 77, 130.

Spores plano-convex (lat), 25 x 33 x 25 p (unacct. same). Lacsura 28 p. Esinc 1.5 p thick, densely rugdate (riigulac dark-brown). Many of the thickenings are short, sometimes appearing like spinules, about 3 p long. Perixie absent.

53. C. heferocnrptts (I31.) Ching (1063; Parlobjig, JIiddlc Andamans; G. S. Srivastara 79516).

Spores 26 X 35 X 30 p. Esinc 2 p thick, not clearly subdivided into sesine and ncsine, smooth. I’erine closely adherent to esine, l~rown, very sparsely spinulose, wrinkled into rugulatc folds pro- truding up to 4 p froin the esine surface. Ridge over lacsura elongated.

74. C. hferritpfcts (Wlld.) Ching (G80; Sahdol, JIadhya I’ratlesh; P. Chandra 95143). Fig. 78.

Spores 24 X 3G X 24 p (unacet. same). Lacsura 20 p. Esine 2.G p thick, not clearly subdivided into sesine ant1 ncsine, smooth. Perine 5 - 631134 Grsnn Pal. rol. 6 : 1

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98 I3. I i . XATAR A S D SASTIIA DEVI

bro\\-nish, closely adherent to esine, smooth, highly wrinkled into sharp-edged, short, rugulatc folds protruding up to 4 p from wine surface. Ridge over Iaesura elongated.

75. C. molliiisculiis (U’all.) Ching (430; Goalpara, Assam; S a y r S I’rem La1 50954). Figs. ‘19, 80.

Spores plano-convex (lat), 22 X 31 X 22 p (unacet. same). Laesura 20 p. Esine 1.6 p thick, light lnown (unacet. same). Sesine nearly as thick as nesine, smooth. Perinc nearly hyaline (unacet. same), adherent to esinc, spinulose, \\-rinkled into elongated, thin, more or’less sparse ridges which in some cases present a rugulate pattern and protrude up to 6 p from the csine surface. Spinules and foldings of the perine more prominent in unacetolysecl spores.

76. C. nioiilnieinensis (I3edd.) Copcl. (379; Tim, Garo Hills, Assam; Kayar B I’rem La1 s.11.). Figs. 82, 83, 140.

Spores plano-convex (lat), 25 x 36 X 26 ,u (unacet. same). Esine 2.2 p thick. Sexinc nearly as thick as ncsine.

77. C. pccrasificrrs (Linn.) Farwell (636; Shillong. Assam; Xayar

Spores 27 X 38 X 30 ,u (unacet. same). Laesura 20 p. Exine 1.6 ,u thick. Sesine nearly as thick as ncsine. smooth. I’crine brown, closely adherent to csine, prominently granulose (some of the granula arc markedly larger than the others and sometimes appearing like blunt spinules) and sparsely folded into faint, short, irregularly conical folds. Fold over lacsiira broad.

49810).

78. C. prociirrens (Nett.) 0. Ktzc. (1 115; Chantlmari, Gauhati, Assam; Xayar L! Prem La1 50973).

(unncet. 23 X 36 x 24 p). Lacsura 22 p, crassimarginate (with thin smooth lips). Esiiie 2.6 p thick. Sesinc nearly as thick as nesine, smooth. I’erine light brown, sparsely spinulosc, closely adherent to the esine, sparsely folded into short, irregular ridges protruding up to G p frorn esine surface. Unncetolysed spores with small, sparse pale green chloroplastitls and one or two pale yello\v oil globules.

Spores 23 x 40 X 28

79. C. siibpibescens (Bl.) Ching (1061; Wimberlyguiij, South Anda- mans; G. S. Srirastara 78762).

Spores plano-convex (lat), 26 x 36 X 26 p. Laesura 24 p. Esine

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SPOI\E >IOI\PIIOLOGT OF I S D I I S P E R S S 99

1.6 p thick. Sesine nearly as thick as nesine. smooth. I’erine light ljrown, closely adherent to esine, faintly granulose, folded into sharp-edged, rugulate ridges.

80. C. iriphyllru (Sw.) Copel. (920; Jorhat, Xssam; P. Chandra 89055). Figs. 81, 141.

Spores elongate-oblong (lat), 26 x 43 x 28 ti (unncet. 24 x 39 x 21 p). Laesura 26 p, its margins incrassate. Esine l . G ,u thick. Sesine nearly as thick as nesine, smooth. Perine dark brown (unacet. same), closely adherent to esine, faintly granulose, highly wrinkled into blunt-edged, rugulate ridges protruding up to 4 p from the esine surface. Ridge over Iaesura elongated.

81. C. urophyllrrs (Wall.) Copel. (589; Pynursla, I<. SC. J. Hills, Assam; Xayar 7i312). Figs. 84, 142.

Spores elliptic (pol), 28 x 42 x 28 p (unacet. 24 x 36 x 25 p). Laesura 24 p. Esine 3.3 p thick. Sesine nearly as thick as‘ nesine, smooth. Perine nearly liyaline (unacet. bro\vnish), loose, faintly and sparsely granulose (unacet. granula more evident), folded into thin, irregular ridges most of which are elongated parallel to the equatorial asis of the spore, and protruding about 8 p from tlie esine surface. Small but prominent, pale-yellow oil globules and dense chloroplasts occur in unacetolysed spores.

19. AMPELOPTERIS Kunze 82. A. prolifera (Retz.) Copel. (664; Satna, Bihar; A. Singh 54885). Figs. 85, 88, 144.

Spores monolete, bilateral, plano- to concaro-conres (lat), ovate (pol), 26 X 47 X 30 p (unacet. 19 X 29 X 18 p). Laesura 20 p, tenui- margin a te.

Esine 2.2 ,LC thick, brown (unacet. same). Sesine nearly as tliicli as nesine, smooth. I’erine light brown (unacet. same), more or less loose, densely spinulose, sparsely folded into short, thin, lobate or sometimes angular ridges protruding u p to 5 p from the esine surface.

20. SPHAEROSTEPHXXOS 6 . Sm. 83. S. polycrtrpa (Bl.) Copel. (1134; Kollem, Goa; 1’. Cliandra 95503). Figs. 85, 86.

Spores monolete, bilateral, plano- to slightly concavo-comes (lat), oblong (pol), 30 X 44 X 34 ,u (unacet. same). Laesura 20 p, tenui- marginate.

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Exilic light brown, 3 p thick, not clearly diflerentiatcd into sesine and ncsinc, densely spinulose (spinules uniformly distributed). I’crinc closely adherent to esinc, nearly indistinguishable csccpt for the fold over the laesura. Unacetolysecl spores with dense, nearly colourless plastids and a few faintly colourctl oil globules.

21. STEGNOGRX;\IX4 131.

S4. S. aspidioides 131. (293; Shillong Peak, Assam; Nayar 50417). Fig. 89.

Spores monolete, hilateral, plano- to slightly concavo-conves (lat), oblong (pol), 25 x 44 X 32 p (unacet. 26 X 38 x 28 p). Laesura 24 p, crassimarginatc.

Esinc 2.2 p thick, light brown (unacet. dark brown), not clearly differentiated into sesine and nesinc, densely spinose. Spines slender, pointed, 4 p long. Pcrine absent.

22. DICTPOCLISE Sloorc

85. D. griffifliii Sloore (517; Pynursla, I<. S J. Hills, Assam; Nayar 77317). Figs. 92, 143.

Spores monolete, hilateral, plano- to faintly concavo-conws (lat), oblong (pol), 38 X 57 x 30 p (unacet. 34 x 54 x 33 p). Laesura 28 p, tcnuimarginate.

Esinc 2.2 p thick, brown (unacet. dark brown). Scsinc slightly thicker than ncsine, tlenscly spinose. Spines dark brown, 6-8 p long, sometimes slightly curved, sharply pointed or blunt. Pcrinc absent. There is a tendency for the spores to split through the laesura into two ralves, even when mounted directly (without acctolysis or centrifuging). Unacctolgscd spores contain dense, pale green plastids.

23. CYSTOPTBRIS nernh.

SG. C. fragilis (L.) Bernh. (1 120; Liddervat, Ihshmir; Janki Prasacl 1007G). Figs. 90, 01.

Spores monolcte, bilateral, plano-convex (lat), elongate-oblong (pol), 29 x 43 x 28 p (unacet. same). Lacsura 24 p, tenuimnrginatc.

Exinc 2.G p thick, deep brown (unacet. same). Sesinc thicltcr than ncsinc, spinosc with the surface between the spines granulosc. Spines deep reddish brown (unacet. same), with a broad h s c , tapering to a pointed apes, sometimes curved and up to G p long. Pcrine absent .

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SPORE MORPIIOLOGT OF I S D I A S FEIISS 101

24. DIPLAZIUM Swartz

S i . L). asperum B1. (1 31; Tiger Shola, I<odail;anal, AIadras; Xayar 45381). Figs. 95, 145.

Spores monolete, hilateral, concavo-conws (lat), 01)long (pol), 35 (30 - 4) x 55 x 36 ,LC (unacet. same). Lacsura 30 p, tenuiniarginate.

Esinc 2.2 p thick, light brown (unacet. same). Sesine slightly thicker than nesine, smooth. I’erine nearly hyalinc (unacet. same), faintly granulose, closely adherent to esinc, either without folds or very sparsely and lightly folded into elongated ridges, ant1 pro- truding up to 6 p from the esine surface. Unacetolysetl spores con- tain a few chloroplasts and a large number of small, nearly uniformly distributed oil globules, two to four of which are conspicuously larger and Yiolet tinged. In some spores there is often a large central, irrcgular oil droplet forniccl by coalescence of the large globules.

88. D. bantamense Ul. (1119; Cherrapunjce, IC. 6 J. Hills, Assam; Xayar 63474).

Spores plano- to concavo-conres (lat), orate (pol), 28 x 42 x 25 p (unacet. same). Laesura 20 p. Esine 1 . G p thick. Perine absent. Unacetolysecl spores with large, dense, prominent plastids.

89. L). bellum Clarke (1 1 1 1; Kodaikanal, Madras; Kayar 44487). Spores ovate (lat), 24 X 40 x 28 p (unacct. same). Laesura 20 p .

k i n e 2 p thick. Sesine nearly as thick as ncsine. I’erine light hrown, loose, smooth, folded into elongated, irregular, thin, more or less sparse ridges (occasionally lobate), mostly elongated parallel to the polar asis of the spore, and protruding tip to 16 p from the esine surface.

90. D. dilafatizrrz B1. (1 108; Cult. S.U.G.). Spores elliptic (lat), 30 X 14 X 30 p . Laesura 23 p. Esine 1.5 p

thick, brown. Sesine nearly as thick as ncsine. I’erine light brown, niore or less loose, sparsely folded into thin sinuous ridges protruding up to G p from the esine surface and mostly parallel to the equatorial axis of the spore.

01. D. escitlentiirii (Retz.) Sw. (1150; Cult. X.U.G.). Figs. 03, 94. Spores 26 (20 - 3) X 46 X 24 p (unacct. 24 X 1 2 X 24 p). Laesura

22 p. Esine brown. Scsiric nearly as thick as nesine, densely rugulate with short irregular rugulae. Perine absent. Unacetolyscd spores with dense, nearly hyaline plastids and one or two large oil globules.

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102 B. I<. S A T A R AXD SASTllh DEW

92. I). j q o n i c u m (Thbg.) Bcdd. (69s; Tiger Shola, Kodaikanal, Madras; Xayar 41463). Figs. 06, 07.

Spores plano-conres (lat), elliptic (pol), 30 x 40 X 32 ,u (unacet. same). Laesura 2G p, its margins incrassate. Esinc 3.3 p thick, brown (unacet. same). Scsinc nearly as thick as ncsinc. I’crine deep brown (unacet. brown). smooth, highly wrinklcd into densely spaced, irregular, short, hlunt ridges appearing like flat papillae and protruding about i ,u from the esine surface.

93. 19. poIgpodioides 131. (513; Clierrapunjee, 11. LFC J. Hills, Assam; Xayar G34iG). Figs. OD, 146.

Spores plano-conves (lat), elongate-ovate (pol), 39 x 59 x 39 p. Laesura 28 p. Esinc 3.3 p thick. Sesine nearly as thick as nesine, faintly granulose. I’crinc pale bro\vn, loose, smooth, folded into a few, long, blunt ridges and protruding up to 12 p from the esine sur- face.

25. XI’HI’RIUAI Itoth.

94. A. fulcrrttmi Uedd. (1003; Pnnchrnarhi, Bihar; Hira La1 33248). Spores monoletc, bilateral, plano- to slightly concaro-convex (lat),

elliptic (pol), 30 X 46 X 32 p (unacet. same). Laesura 27 p, faintly crassimarginate.

Esine brown (unacet. same), 2 p thick. Sesine nearly as thick as nesine, smooth. Perine brown . (unacet. same), loose, sparsely spinulose, lightly folded into a few, thin irregular ridges protruding up to 8 p from the esine surface. Unacetolysed spores with a few small chloroplasts; spinules not apparent; foltlings of perine more crirl en t . 90. A. filix-fen~incr (L.) Roth rar. (15; Gauhati, Assam; Ram I’hcr 13541).

Spores 2G X 37 x 23 p. Laesura 24 p, tenuimarginatc. Esine 1.0 p thick, not clearly difierentiated into scsine and nesine. Perinc light brown, irregularly granulose, protruding up to 5 p from csine surfacc.

96. A. Jinibriafurn (Wall.) JIoore (1 10’7; Kainital, Uttar Pradesh; S. I<. Jain, 9682). Fig. 101.

Spores obIong (pol), 21 x 32 X 21 p. Laesura 1G p, tenuimarginate. Esine light brown, 1 p thick. Perinc light hrown, adherent to the esinc, faintly granulose, protruding up to 4 p from csinc surface.

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S P O R E JIOIIPIIOLOGY O F I S D I A S PERSS 103

97. A. ~liolienackeriarlomi (Iize.) Jloore (1 1 1 5; Jaipur, Rajasthan; G. S. Srivastava 38619).

Spores plano-convex (lat), oblong (pol), 30 X 42 X 32 p (unacet. same). Lacsura 24 p, tenuimarginate. Esinc 2.2 p thick. Pcrinc light brown (unacet. same), smooth, highly wrinkled into thin, angular, elongated ridges (some of which appear lobate) protruding up to G p from esine surface. Unacetolysed spores with dense granular contents.

98. d. p e c f i m f i z m (Wall.) Pr. (734; Simla Hills, I’anjab; T. Jlukerjee, 15662). Fig. 100.

Spores plano-convex (lat), ovate (pol), 28 X 40 X 30 p (unacet. 24 x 36 x 24 p). Laesura 22 p, tcnuimarginate. Esine 1 . G p thick, light brown (unacet. darker). Sesine thinner than nesine. Perinc light hrown (unacet. deep brown). more or less loose, smooth, highly folded into thick, irregular, blunt folds some of which appear lobatd, and protruding up to 5 p from esine surface.

99. A. procerimi (Wall.) Jlilde (392; Tura, Garo Hills, Assam; Xayar c!? Prem La1 50916). Fig. 103.

Spores plano-convex (lat), oblong (pol), 21 x 32 x 21 p , Laesura 22 p, its margins incrassate. ISsine 2.6 p thick, darkish brown. Sesinc thinner than nesine, faintly granulose. Perinc nearly hyaline, adherent to esine, smooth, sparsely folded into elongated, thin, sparsely sinuous ridges protruding up to 6 p from the esine surface.

100. A. solenopferis (Kze.) Xloore (1 113; Girnar, Snurashtra, Gujernt; Janki Prasad 14086). Fig. 102.

Spores oblong (lat), 26 x 39 X 28 p. Laesura 20 p, its margins incrassate. Esinc 1.6 p thick. Perinc dark hrown, more or less adherent to csine, smooth, wrinkled into thin, irregular, short ridges (many of which are parallel to the polar asis of the spore) protruding up to 6 p from the esine surface.

101. A. spcfcrBife (Vrall.) C. Chr. (553; Iiashinagar, Ileoria, Uttar I’radesli; A. Sing11 63515). Fig. 08.

Spores plano-convex (lat), oblong (pol), 23 X 32 x ‘13 p (unacet. same). Laesura 20 p, tenuimarginatc. Esinc 1.3 p thick, not clearly diflerentiated into sesine and nesine, faintly and densely granulose. I’crine faintly granulose.

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104 13. Ii . SXTAR A S D S.\STIIA DE\'I

102. A. tomentosum (121.) llildc (GOG; Bihar; J. G. Srirastara 53632). Spores concavo-convex (lat), elongate-oblong (pol), 30 (34 - 1) x

4G x 3G p (unacct. 24 (2G - 2) X 43 x 28 11). Laesura 20 p, tenuiniar- ginatc. Esine 2.2 ,u thick. Scsine thicker than nesine, faintly granu- lose. Pcrine absent. Unacetolysecl spores with small, sparse, nearly colourless plastids and one or two faintly yellow oil globules.

111. Discussion

The Aspidiaceae, -as conceived by Copeland (194i), is a large family of homosporous lcptosporangiate ferns, believed to have originated from estinct ancestors in the Antarctic region. It is evi- dently a n ancient group with no near relatives among modern ferns. Four or five major groups of genera, all probably coordinate in crolution, arc recognized: the Polysticlircnt-groLip, the Dryopteris- group, the Tecfnria-group, the Lastrea-Cyclosoriis (Thelyp1eris)- group and the Athgriitm-group. I3esides, there are a few primitive genera of doubtful affinities, .for instance the onoclcoitl ferns (Mat- terzccia) and Pernnema and its allies. Most pteridologists regard Copeland's Aspidiaceae as an unnatiiral assemblage: the dilferent groups of genera are often treated as separate families (Ching 1940, Holttum 1947, Alston 195G, Pichi Sermolli 19GO). A dennstacdtioid (Holttum 1047) or cyatheoicl (Christensen 1938, Ching 1040, I3o\\-er 1023-28) affinity is often suggested for many groups of these ferns, and some, like the dthgrium-group, arc sometimes regarded as having an affinity with the Asplcniaceae (Ching 1040, Pichi-Sermolli 1960).

The Aspidiaceae is charactcrised by the possession of bilateral spores, with generally smooth esinc and a well developed although often loose perinc. Nearly all genera commonly regarded as primitive, possess a granulosc perine. In some genera there is a tendency towards reduction of the perine. In some cases there is apparently no perine at all. In the Cyathcaceae and the Dennstaedtiaceac, cliaracteriscd by the possession of typical trilcte spores, there is. 1il;ewise no perine. A direct derivation of the aspidioid ferns from

Figs. 1-13. ( E , equatorial diameter; n, nerine; P, polar axis; p , perine; s, sesine. All figures, unless specified, are lateral views). - 1, Explanatory drawing (cf. test, p. 81) of a bilateral spore in lateral view. - 2, lllalteuccia orientalis. - 3, Diacalpe aspidioides. - 4 , Peranerna cyatheoides. - 5 , ttcrophorus slipellalus. - 6 , Same, polar view. - 7, ZIgpodemalium crenalurn. - 8, Lilhostegia foeniculacea. - 9, Same, polar Y i e x . - 10, Polysfichurn acanthophyllurn. - 11, Same, esine stratification. - 12, Polusliehum

squarrosum, polar view. - 13, P . squarrosum.

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SPORE 1IORPIIOLOGY O F I S D I A S FERSS 105

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Figs. 14-29. - 14, Polysfichum lobalum. - 15, 1'. aculealum, mine stratification. - 16, P. aculealum, surface view of the perine, showing ornamentation pattern. - 17, P. acrrleafum. - 18, P, slimulans. - 19, Cyrlomiirm hookerianum. - 20, Exine stratification of the same. - 21, C. falcafum. - 22, Xurnoitra carvifolia. - 23, Bolbifis presliana, .ornamentation pattern of the perine. - 24, Uolbilis presliana. -' 25, B. coslala. - 26, B . semicordda. - 27, Same, polar view. - 28, Esiric stratification of

the saine. - 29, U . crispatuln.

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Figs. 30-41. - 30, Bolbilis virens. - 31, B. heleroclila. - 32, B. subcrcnala. - 33, B. sicbsimplex. - 31, Egenollia sinensis. - 35, Same, polar view. - 36, E . sinensis, polar view of a spore with forked laesura. - 37, E. appendiculala. - 38, E . asplenilolia. - 39, Omanrentation pattern of the pcrine of the same. - 40, E . vivipara. - 41, Ela-

phoglossirm con farme.

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55

Figs. 42-55. - 42, Dryopteris lilix-mas. - 43, D. barbigera. - $4, D. hiriipes. - 43, Cleriifis hendersonii. - 46, Teclaria ciccclaria, polar view. - 47, T. fusipes. - 48 , Exilic stratification of the same. - 49, T. deoera. - 50, Polar view of the same. - 51, T. polymorpha, polar view. - 52, T . polymorpha. - 53, Esine stratification of the same. - 51, T. macrodonla, esine stratification. - 55, T. macrodonfa.

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62

Figs. 5G-71. - 56, Teclaria semibipinnala. - 57, Esine stratification of the sntnc. - 58, T. simonsii, esinc stratification. - 59, T. simonsii. - G O , T . subconfluens. - G1, Esinc stratification of the same. - 62, T. uarioloso. - 63, Qiiercifilir reilanica. - G1, Laslrerc calcarala. - G5, Polar view of the same. - G G , L. crassifolio. - 67, I.. falciloba, esine stratification. - 68, L. /nlciloba. - 69, L. gracilescens. - 70, Esinc stratification

of the same. - 71, L. phegopferis.

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87 Figs. 72-89. - 72, Laslrea puluinulifera. - 73, L. pgrrhorfiachis. - 74, Polar view of the same. - 75, Cyclosorirs crislala. - 7G, C. didymosora. - 75, Polar view of the same. - 58, C. inlerrupfirs. - 79, C. mollircscultrs. - 80, Polar viev of the same. - 81, C. lriphyllus. - 82, C. moulmeinensis. esine stratification. - 83, C. moulmeinensis. - 81, C . urophyllus. - 85, Sphaeroslephanos polycarpa. - 86, Esine stratification of thc same. - 87, dmpelopteris proli/era. - 88, Exine stratification of the same. -

89, Slegnogramma aspidioides.

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to I 101

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.Figs. 104-1 16. - 104, Peranemn cgnlheoicles. - 105, Diacalpe ospidioicles. - lOG, Lilhoslegia loenictclacea. - 107, dcrophorus slipellalus. - 108, I~olyslichum biarislalum. - 109, P . speciosum. - 110, I>. auriculalom. - 111, Cgrlomictm hoolierianum. - l l l n , Rrrmohra carvifolia. - 112, Uolbilis coslaln. - 113, B. cosfafa, polar view showing forked lacsura. - 114, U . crispaluln. - 115, L). subsimples. - 1113, n. presliana.

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Pigs. 117-128. - 117, Dolbilis subcrenala. - 118, Egenolfia vivipara. - 119, E. sinensis. - 120, nolbilis semicordala. - 121, Egenolfia appendiciilnla. - 122, Elaphoglossum conforme. - 123, Drgopleris filit-mas. - 121, D. hirlipes. - 126, Laslrea falciloba. - 126, Drgopleris cocltleala. - 127, Laslrea pulvinulilera. - 128, L. pgrrhorliachis. 8-G31134 G r a m Pal. vol. 5 : 1

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114

Figs. 129-137. - 129, Clenilis hendersonii. - 130, Teclaria deitera. - 131, T. seniibi- pinnuta. - 132, T. subcon/luens. - 133, 3'. fuscipes. - 134, 3'. polpnorphn. - 136,

2'. ciculario. - 136, T. variolosa. - 137, I~olgs1icIium sguarrosum.

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SPOI\E SlORPlIOLOGT O F I S D I h S FEIWS 115

Figs. 136-l4G. - 136, Cyclosorizs crislula. - 139, C . tlidymosorn. - 140, C . moiilmeinen- s is . - 141, C . lriphylliis. - 142, C . urophyllus. - 113, Diclyocline gri//ithii. - 144, Anlpelopleris prolileru. - 145, Diplazium asperurn. - 14G, D. polypodioides.

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tlie Cyatlieaceac or the Dcnnstaedtiaccae is, however, open to doubt. Copeland’s contcntion that they are evolved from extinct ancestors appears to suit tlic facts best.

Regarding affinities, the genus PolrJstichiim is regarded by Copc- land (194’7) as derived indcpenclently from estinct ancestors, while others (Ching 1940, Holttuni 1947) postulate a dryopteroid affinity and relationship to peranenintoid ferns (Bower 1023-28). The characteristic nature of tlic pcrine in Z’olysfichrim docs not lent1 support to tlie suggestion of a relationship to either Permenin and Dicictilpe, or to the clryopteroid ferns. X loose perinc wrinkled into small, lieinisplierical, crowded folcls is interpreted as an indication of “primitivcncss” in Z’olgstichuni. In Z’. lobatuni, 1’. sqii(rrrosiiiii and Z’. sliniirlans among Indian species and Z’. crespidosoriini, 1’. p o l p blepliciriiiii, Z’. tripteriis and Z’. tsiis-siiiieiisc among Japanese species (Sayar & Dcvi 19G3) tlic folds of the perinc are lobate. Since these species are regarded as more or less “advanced”, the condition of tlie pcrine is prcsuma1)ly a clcrired one. In Cyrfoniirrni, a polysticlioid derivative, the perinc is somew-hat similar to that of these “admncetl” species of Polyslichrmi. But in C . hookerinnrrni, which resembles Polrpfichrcni in many morphological features and, therefore, seems to be more close to l’olystichum than otlicr species of Cgrtoniiiiiii. the perine is fol(lec1 into hemispherical, cro\vdecl folds-much as in the more primitive species of Polystichiitn. In C. cnryotiderini and C . frrlccitiim, just as in some species of Dryopteris, the lobes of the pcrinc have a tendency to become convolute or tulxrculate. Among other ferns of “po1ysticlioid” affinity, the spores of Rrrniolirn (Byrsopteris) deviate from those of Polysticlirmi: the perine is different in its pattern of folding.

Bolbitis and Egenolfia arc sometimes referred to the Lomariopsi- daceac (Holttuni 194i). Bolbilis is regartled as older than Egenolficl (Copeland 1047). In the more “primitive” species of Bolbitis ( R . prcsliancr, B . seniicordata) the pcrinc is prominently granulosc and folded into subconical folds, resembling tlie common type of perine in Polysticliirni. In inorc “adranced” species, the pcrine is smootli and folded into lobate ridges, as in tlie supposedly more “adwnccd” species of I’olysticliirm. Egcnolfia dilTers from Bolbitis in its characteristic reticulate ornamentation of tlic pcrine. As to affinities, Copeland (1947) mentions tlie possible relationship of tlie group to tlic Polysticliiini-Riiniolirrr-6roup. As far as Bolbitis is concerned, spore morphology appears to support this contention.

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S P O R E JIOHPIIOLOGY O F I S D I A S FEltSS 117

~iolttuni (1 9-i.i-1 040) includes Elaphoglossurn in the Loniariopsi- daccac, hut other botanists regard it distinct enough to constitute a family by itself (Christensen 1938, Cliing 1940). The spore morpho- logy of Elriphoglossiini conforiiie does not indicate any close affinity to Egenolfici or Bolbitis, nor docs it support a dryoptcritloid relation- ship as suggested hy Copelancl (1947).

The tectarioitl ferns (Ctenitis, Tectaricc, @iiercifili.r) are lxlic\-ed to have had a coninion ancestry with Dryopteris: according to Cope- land (101i) there is a n apparent affinity to Riimolira. In the genus Tectaria, T. clevcscr and T. fiiscipes etc. have a loosely folded, spinu- lose perinc antl a smooth esine, T. polymorpha a loose, smooth pcrinc antl an ornamented csinc, while in T. helerosorcr the perine is spinose, closely adherent to the granulosc esinc. It thus appears that there are two separate. lines of evolution among the species stutlictl: (1) from those with smooth csine antl smooth pcrinc, as T. vciriolosn (resembling the probable parent genus Ctenitis), through T. polymiorphci with smootli loose perinc antl spinulose csine; (2) from T. dei)e.rct and T. fiiscipes with loose spindose pcrine, through T . siniorisii with less prominent but spinulose perinc, to T . semibipinnatci and T . helerosoru u-ith pcrine clyscly adherent to csinc and prominently spinosc. From a spore morphological point of view, @tierci/7li.r (wilh smooth esine antl loose smooth perine) appcars to he dcrivctl from a group of species in Tectcirici represented hy T. unriolosn.

The Thclypteridaceac consists of Lasfrea (Tlielypteris), Cyclosorirs antl their derivatives. Lnslrea and Cgclosoriis are so intimately related that many ptcritlologists merge the two. They are hoth very large, characterized hy faintly ornamented esinc and perinc. There is a m d i e d tcntlency-morc evident in Cyclosoriis than in Lasfrea- for reduction antl loss of the pcrine. The tendency of the pcrinc to have a crest-like elongated fold at the prosinial pole (opposite the ladsura) is characteristic. The iinmcdiatc derivatives of Cyclosoriis (Sphaerosfeplinnos, Sfegnogmmmcr and Dictgodine) are devoid of pcrinc (or nearly devoid of it), but possess granulosc or spinulose csinc, as, e.g. in C. nioiilmienensis, C. crinipes, and Lcistreci fcrlcilobci. Ampclopferis, though derived from Cyclosoriis, difkrs by having a distinct, spinulosc pcrine and a smooth esinc (esamples: C. molliris- ciiliis antl C . lieterocarpiu).

Athyriiini and Diplariiim are old ferns, often regarded as evolved independently from extinct ancestors and deroid of any near rela-

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118 I3. I<. S . iTAR .\SD S.iSTI1.i DEYI

tires (Copeland 1947). As in the thelypteridoid genera, there is a marked tendency for the reduction and loss of the perine as \veil as for the clevelopnient of ornamentations on the csine. Many Japanese species of dfhyrizrni (Nayar & Dcri 19G3) and Indian species of Diplaziizni exhibit this tcndency. Another line of advance- ment in these ferns appears to be towards the clevelopment of a n ornamented peritie as in dthyrizim falcatrzni.

Among the more primitive genera of the Aspidiaceae, Jlafteizccicl possesses spores which are unusually large in the family. The perine is loose and both perine and esine are smooth. The spore morphology of -11. oriezitalis does not appear to support the suggestion (see Nayar 1961) of its affinity to either the Cyatheaceae (with trilete spores) as suggested by I3o\ver (192G-%), Christensen (1938) and Ching (1940), or to Percznenicz and Uiacdpe as suggested by Copeland (1947). Spores of some blcchnoid ferns (Xayar 19G2) do. ho\verer, resemble those of Jlaffeuccia (except in size). Diclcalpe, I’eranenia, Acrophorzis and Lifhostegia are commonly regarded as closely related ferns, and their spore morphology supports this view. Copeland (1947) and Ching (1940) consider Lifliosfegia as heing related to Polysfichizni, while Acrophorzzs “fits with the Darallioid ferns better than with any otlrers” (Ching 1940, p. 220). The spores of these genera differ markedly in their morphology from those of the Poly- stichizni-Rrrniohrcl-group of ferns.

From the present study it secms evident that acetolysis does not necessarily result in an increase of size of the spores in all cases. Some genera, e.g. dfhyr izm, Cyrfominni, Dipla:ium and Lasfrea, do not show any significant size changes as a result of acetolysis. Alost species of Cyclosortzs, Polysficliimi and Tectaria behave similarly. Also, in those spores which increase in size on acetolgsis, the per- centage of increase differs widely in the different species, some like Anipelopteris prolifera eshibiting about Gf! ‘$6 increase (in El), Urgop- feris hirfipes 50 % (El), Egenolfia asplenifolicl and Elaphoglosstzni con fornie 43 % (El), Diacalpe, Bolbitis cosfata and Dryopteris filix- nim 35 % (El), etc. For any given species, thc increase in €lie dif- ferent directions (P, El, E,) is not proportionate. In Polgsfichum discref tzm and Sfegnogramma cispidioides the P is unaffected while El and E2 increase by 25-30 % and 1G-23 %. In Lithostegia foeni- ciilacea and Runiohra aristafa the E2 is unaffected while the P and El increase by 18-25 % and 14-24 % respectirely. IR Cyclosorizs tri- phyllizs the P and E, are unaffected while El increases by 25 %. In

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S P O R E JIOI1PIIOI.OGI O F I S D I z \ S FEllSS 119

Elnphoglossrrni confornie the P increases by 8 % while the El and E, increase 1)y 42 ant1 18 % respectively. In Arnpelopferis polifera the 1’ increases hy 36 x, El G2 YL and E2 GG %. It thus appears tliat tlic iiicreasc in size of spores on acctolysis is a specific character deter- mined by the structure of the spore-wall in cncli species.

IV. Summary

Spore morphology of 102 ferns from 25 genera of the Aspitliaceac is tlcscribetl. The spores are monolete, hilateral, piano: to more or less concaro-convex, varying from 20 x 30 x 22 p to 45 X GO x 52 p. Tlie lacsura is usually tcnuimarginatc. The esinc is mostly smooth (niorc rarely granulose, spinulosc, or spinosc). As a rule, tlie spores have a pcrinc, but in some groups of genera the pcrinc is rednced and eventually lost. The perine is commonly granulose and charac- teristically folded, in sonic cases it is smooth or spinulosc. Acetolysis results in increase in size of spores in some species; tlic extent of increase in each plane appears to he a specific character. The significance of some of the morphological features of the spore in phylogenetic studies of the various lines of evolution in the Aspi- tliaccae is discussed.

Acknowledgements

The keen interest evinced in this work by Prof. I<. N. ICaul, Director, National Botanic Gardens, is gratefully acknowledged. U’e are t1ianl;ful to Prof. G. Erdtnian for kindly going through tlie MS. l‘o Jlr. 1’. C. Roy we are thankful for his help in the preparation of the illustrations. Tlic junior author is grateful to the Council of Scientific and Industrial Research (India) for the award of a fello\vship to her, which made this study possible.

Ptcridology Laboratory, National Botanic Gardens, Lucknow

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