This article was downloaded by: [University of Georgia]On: 17 December 2014, At: 21:17Publisher: 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 ofthe Pteridaceae III. TheDicksonioid, Dennstaedtioidand Lindsayoid FernsB. K. Nayar a & Santha Devi aa National Botanic Gardens , LucknowPublished online: 01 Sep 2009.

To cite this article: B. K. Nayar & Santha Devi (1968) Spore Morphology of thePteridaceae III. The Dicksonioid, Dennstaedtioid and Lindsayoid Ferns, GranaPalynologica, 8:1, 185-203, DOI: 10.1080/00173136809427464

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SPORE AIORPHOLOG1’ OF THE I’TEIUDACME 111. THE DICKSONIOID, DENNSTAEDTIOTD

AND LINDSAYOID FERNS

J I S receiiletl I7.51 I .6 ;

Introdiiction

In tn-o prcvioiis coiiimuriicntioiis (Sayar and Dcvi I O G G , I O G i ) wc Iiavc tlcscribctl the s p r c morphology of tlic ptcroitl arid tlic gyiiino- graniiiioid fcrns, togetlicr constituting tlic major part of the I’tcri- claccac as prcscntctl by Copc1:intl (1947). ‘I’lic tliclxonioid, tlcnn- stactltioid nntl liiidsayoitl fcrns tlcnlt with in this p p c r arc listed 1)y Copelnntl as gcncrn no. 2-29 of his I’tcritlaccac. ‘I’licsc groiips o f fcrns h a w for a long tinic hccn problcmatic from tlic taxonomic and phylctic points of view. So tiyo tasoiioiiiists scclii to agrcc on thc rclationsliip of the various gcncra constituting thcsc groups. ‘I’lic estcnt of t1is:igrccnictit that currcnkly csists is well rcllcctctl in tlic various \vays tlicsc gCliCra a rc trcatctl in tlic tliITcrcnt systems of classification prcscntctl in rcccnt days (Cliristcnscn 1938, Ching 1940 , Copclantl 194i, IIolttum 1 W i , Pichi-Scrmolli 1958, 1059). -hiorig morlcrn ptcritlologists, Cliing (1940) groups tlicsc gciicra of fcriis untlcr a niim1)cr of scparatc familics-Culcitaccac, Dcnn- stacdtiaccnc, Hypolcpidaccac, Lintlsayaccnc-all of u-liicli in his view arc distinct fro111 thc ptcroitl nncl gyiiiiiogr:iiiiiiioitt fcrns. Piclii-Scrniolli (1 958, 1950) supports this vicn-, rcfcrring thcsc fcriis to the Ortlcr I)icliso~iinlcs (familics I)iclisoniacc:ic, 1)cnnstacd- tiaccac and Lintlsacaccac) distinct from thc Ortlcr l’teritlalcs which incliidcs the ptcroitl ant1 the gyninogramnioitl ferns. Cliristcnscn (1038) trcats thc tlicksonioitl group of genera as a family scparatc from his ‘I’olypotliaccac’ which incliidcs tlic ptcroitl, tlic gyniiio- gramrnoitl, the lintkiyoitl and thc dcnnstactltioid fcrns. Holtturn

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(1 947) slipports this view in rccognising tlic Dicksoniaccac. regards tlicni as rc1:itctl to tlic family of trcc ferns, tlic Cyatlicaccac. Howcver, lic csclutlcs the gyinnogramnoid genera froin thc others, grouping tlicm under his Xdiantaccac. Also, thc ptcroitl antl lin(1- snyoid fcrns arc conceivcd as endproducts of divcrgcnt cvolutio1l from ancestral typcs similar to tlic dcnnstncrltioid fcrns. Thcy arc, thercforc, includetl in tlic large family Dcnnstactltiaccac (ivliicll inclu(1cs also tlic davallioiti, dryopteri(Ioit1 an(1 asplenioii ferns). Tlic dcnnstacdtioid antl lindsayoid ferns arc gronpcd in t\vo suh- familics of equal r ank as tlic Asplcnioitlcac. Davallioidcac, ctc., which arc rcfcrrctl to indcpcndcnt familics by most ptcridologists. Recently, based on cstensivc cytological cvidcncc, SIanton (1958) advocatetl the tasorioinic separation of the diclisonioitl, t1cnnst:ictl- tioid and lindsayoid fcrns from thc otlicr gcncrn of Copeland’s Ptcrirlaccac. Shc contends that for these fcriis “it is probablc that rccogriition of inorc than onc family coiiltl l x s\lggestcd, ant1 may 1)c ncccssary” (,\Innton 1‘358. I). SG). Alelira (10G1), also on cgto- logi cal ex-i d cnc c, r ccoiiiiii en cls that Hist iopl eris, H ypolepis, Pa esiri and Ptcridiiim, bc rcfcrrctl to a scparatc faiiiily (IIypolcpidnccnc) pliy1ctic:illy distinct from tlic Dicksoniaccac and Dcnnstaetltiacc:ic: also, he docs not consider the latter two f:iniilies as closely inter- related. Xl~raliani et nl. (10G2) regards this vie\\, as “unconvincing and based on incorrect appreciation of cytological data”. Tliey suggest iristcatl that thcsc genera, csccpt tlic tlicksonioitl fcrns (\vIiicIi constitiitc the Diclisoniaccac) should 1)c rcfcrrctl to one family. ‘I’his family, provisionally callctl thc Dcnnstacdtinccac, is in their opinion rclatcd to the 1)icksoniaccac on the onc hand and to the Hy1iic~iol)Iiyllnccac and Davalliaccac on the other.

‘I’his v-itlc variation among tlic difTercnt ptcritlologists in tlic inter- pretation of thc phylogeny and rclationship of tliesc fcrns cvidcntly indicates thc nccd f6r more morphological information regarding these gcncra. As rcmarkccl by Manton “tlicsc gcncra arc csactly the oncs which tend to hc thinly scnttcrcd, difficult to grow or in other ways inacccssihlc, so that accumulation of sufficient data cannot bc espcctcd to bc otlicr than slo\v” (Manton 1’358, p. SG). It is hoped that the prcscnt account of spore morphology will contribute to n I)cttcr understanding o f thcsc much dc1,atcd gcncra of ferns.

Ncarlg .50 pcr ccnt of tlic specics incluclccl in this study arc Indian. thc material for study collcctcd inninly from Assam and tlic Wcstcrn Ghats. ;\l:itcrial of tlic other species has 1)ccii obtninetl from various

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S 1’0 II 1: .\I 0 II I ’ l l 0 1.0 (il’ 0 I: TI 11: I’T 1; III D.\CEA E. 111 187

p r t s of tlic \vorld from tlic clifTerent agcncics mcntioncd in tlic text. ‘I’hc tccliniquc of s tudy follo\vctl and that of tlie tlcscriptions (in- c luding tlic terminology nsctl) arc t h e same as in carlicr cornrnunica- tioiis of this scrics (Nayar and Devi 19G4 a-c, 1965). The genera arc arrangctl in tlic saiiic \yay as i n Copclantl’s “Gciiera Filiciiin” (19-!7). The species iindcr each genus :ire listctl in alpIia1)ctical ortlcr. I)ctailctl description is provided only for tlie first spccics untlcr cacli genus. For subscqucnt spccics, only tliosc features arc incntioncd \vliich tliflcr from those of the first species. Herliariuni rccords of all tlic Indian species arc maintained i n t h e Herbarium of the Xntiorial Botanic Gardens, Liicknow. ;\laterial obtninctl from otlicr licrbnria arc spccifictl, thc international abbrcviations of the iiaiiies of the 1icrl)aria bcing appcndctl to tlic statcmcnt on tlic soiircc o f tlie matcrial. ?‘he a1)l)rcviations “ncct.” and “iinacct.” starid for “acctolyscd sporcs” ant1 “unacctolysctl spores” rcspcctiwly. All illustrations, iinlcss spccifictl otlicr\vise, arc rcprotliiccd at 750 tiiiics tlie actual s i x , csccpt tlic latcrnl vic\vs of tctralicdral sporcs arid tlic prosimal yicu-s o f 1)iI:ikral spores which arc rcprotluccd a t ,500 tiiiics the actual size. ‘I’lic tcst-figures arc cariicra lucitla drawings. l’licy do not ncccssarily tlcpict a spore of tlic average size nicntionctl in the tcxl.

Descriptions I) I C I< S 0 S I A L’ H cri t i cr

1. L). trrilarclicci 1 ~ 1 ) . (1526; Escli., School 1)ioI. Sci., Univ. Sydney). - Fig. 9.

Spores trilctc, tctralicdral, 46 x GO / I ; range 44-18 x 56-61 / I . Ainb tri:ingular with almost straight sides and broadly rouiitlcd coriicrs. Prosinial facc of spore flat to sliglitly convex; distal facc Iiemispliericnl. Lacsura arins 22 11 long, their outer margins thickened into prominent lips, about 4 / i

broad but distinctly tapering towards the t ip of the arms. Esinc light ycllowisli brown, 2.5 / t thick. Scsinc iicarly as thick as ncsine,

granulose all over and, in addition, arcolntc on the distal facc. Areo1:ic polygonnl, 10-12 / i across, 3 / c Iiigli, separated by sliarply tlcfiiicd grooves about 2 / I broad.

111 n prosimal view of the sporc two sharply defined outlines, one inside the otlicr, arc sccn. ?‘his is bec:iuse the prosinial face of the sporc is narrower than the distal facc and the two faces arc sharply delimited. Thus, wlicn the spore is seen from the proximal pole, the outline of the distal li:ilf is sccn as :i triangle cncircling the triangular outline of the proximal half. This gives a false appe:irance of tlic spore possessing an equatorial collar- like ritlgc as found in Pleris.

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2. 11. /ibrosrr Col. (1G18; X. Xmcrica; Xmer. Fern S O C . no. 274). - Fig. 1.

Sporcs 40 x51 11; range 36-48 x48-56 11. Amb with nearly straiglit to slightly coiicave sides and broad, rather triiiicatc coriiers. Lacsurn arlllS IS 11 long, tlicir outer margins faintly thickened. Esiiic dull browii, 4 11 tIiick. Scsinc thicker tliaii tiesiiic and ornamented in the saiiic \\'ay ;is i l l 1). anfrtrcficci, bu t witli iiiorc prominent granulations.

Uiiacct. tiearly of tlie saiiie size as the acet., grccnisli-ycllo\v. Esiile nearly Iiyaliiie. I'lastitls large, densely crowded.

3. 11. sqirrrrrosri (Forst.) SW. (1804; X. America; Atncr. F e r n Sol:. no. 2iS). - Fig. 2.

Sporcs 5G x SS / I , with little variation in size. Aiiib triangular witli ilcarly straight to slightly coiicave sides. 1,acsura a r m 32 11 long; lips faint, smootil and nearly uniform tlirouglioiit the length of the arms. Esinc nearly Iiyaliile to pale brown, 4 p thick. Scsirie niiicli thicker than ticsine, proiiiincntly granulosc all over, and i n addition faintly uiidulating on tlic prosinial face and with irrcgiilar areolae (8-12 11 across, 3 p high) 011 the distal face.

Unacct. iicarly of the sanie size as acet. Esiiie almost Iiyaliiie, less tl1;lIl 2 11 thick. The sporcs contain 11i:itiy large, crowled, light-grccn plastirls atid many large, deep-gcllowisli oil globules. A notable feature of the spore of D. squctrrosci is that tlie exilic is liable to dissolve coiiipletcly i f Iicatctl for more than a miiiutc in tlic acetolysis iiiisturc. IVIicn spores are mounted i n tlic acetolysis iiiisturc at SO-90% atid observed unrlcr tlic iiiicroscopc tlic csiric is seen to swAl gradually, becoming soft. 1;inally i t dissolves allnost coniplctcly. On tlissolving a few sniall gas bubbles arc produced.

c I 13 O'I'I UJ 1 Iin u 1 f 11 ss

4. C. bnronielz (Lirin.) J. Sin. (G71; cull., L loyd hot. Gartl., D:irjccling, India). - Figs. 4, 12.

Sporcs triletc, 4 3 x 70 11; range 40-48 x G 4 - i G 11. Amb triaiigu1:ir with straight to concave sides ant1 iiarrow corners. I'rosiiiial face of spore llat to slightly concave; distal subconical. 1,acsiir:i arms 28 long, tlicir inner iiiargiiis thickciictl.

Esinc light brown, 3 11 thick. Sesiiie much thicker than ricsiiie, rather iintlulate on the prosimal face and tribcrculate on the distal. The two faces arc scparatctl by a11 almost uniform, equatorial, collar-like ridge protruding about 6 11 from the general surface of tlic csinc. Tubercles 4 p high, 10 11

broad, cloiigatctl parallel to tlie collar arid often coalcscctl to form uiidiilating ridges.

Unncct. 3G x 58 / I , iiearly hyalinc, and containing many large, crowled plastids and a few small, riearly liyalinc oil globules.

5. C . glcciiciim (Sm.) IIool;. & Xrn. (1572; cult., Hot. Gnrtl., Uiiiv. Caiii1)ritlge. ICngland). - Fig. 11.

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Spores 38 x 52 p; range 34-40 x 48-60 p. Ainb triangular witli cotivcs sides and broad corners, to subcircular. Prosirnnl face coIives. Laesur:t arnis 22 11 long, tlieir inner margins tliickencd into lips 8 / i Iwoad :ind sharply t:ipcring towards tips. Es ine darkish brown, 4 11 tliick, densely grnndose.

In ninny cases one of tlie Iacsurn arms is mucli longer tlian tlie otlicrs. Tlic corner of tlic ninb to wliicli tlie longer arm points, is csccssively pro- t r ii (led.

Unacet. 30 x45 p , lighter in coloiir than tlic acet. Contents maskctl.

6. C. schietlei Schlcclit. & Cham. (032 cult.. Ilot. Gnrd., Cologne, Gcrtiiany).

Sporcs 4 G xG0 / I . Laesura arms 22 / I long. Izsine yellowisli-brown, 4.5 / I

thick, vcry faintly granulosc. X thin, outer, skin-like lager licaring the ornnmciitation (? perinc) oftcn cracks into sniall pieccs ant1 sometimes peek off on ncetolysis.

CULCI'I'A I'rcsl 7. C. tlrrbicr (11. Ilr.) JIason (1534; Eseh., School biol. Sci., Unir.

Spores trilcte, 38 x 4 G 11; range 3 6 4 0 x40-52 p. Amb triangular to sub- circular. I'rosimal face of spore convex; distal face hcrnispIierica1. Lnesura arms 14 11 long, their margins untliickeiicd.

Es inc light ycllowisli-brown, 3 11 thick, not clearly differentiated into sesirtc aiicl nesine, densely nrcolnte. Arcoles irregular in outline, varying much i n size (usually 2-8 14 across and 3 11 Iiigli).

0 IZ'I'H I 0 I T I5 It I S Co 1) eln lit1

8. 0. k i n g i i (I3cdd.) Hollt. (682; Shangila Reserve, Quilon, Kerala, India; I3. I(. Kayar 44487).

Sporcs trilctc, 28(30-2) x 40(43-3) 11, with little variation in sizc. Amb triangular with concave sides and broadly rounded corners. Prosimal face concave with protruding, angular pole; distal face subconical. Laesura arnis 10 11 long, with the outer margins tliickencd into faint smooth lips about 3 14 bronc1 on citlicr side of laesurn and tapering towards tlie tips.

Es inc nearly liyaline with an yellowish tinge, 2 / c thick. Scsinc nearly as thick as nesine, smootli.

DEXXSTAED'I'IA 15ernh. 0. I ) . cipperidicultrlti (Wall.) J. Sm. var. elwesi i I3edtl. (652; Darjcc-

ling, India; P. Chantlra 60407). - Figs. G, 14.

Spores trilete, 30 x40 p, with little variatioii in size. Anib triangular (with slightly coiicave to conves sides) to subcircular. Prosimal face of spore coiivcs, with protruding pole; distal face liemispherical. Lacsura arms 16 11 long, their margins unthickcncd.

Sytlneg). - Figs. 3 , 10.

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100 13. I<. S A Y A I I AX'D SAXTIIA D E V I

Esiiic brown, 3.5 /L thick. Sesiiic iicarly as tliick as Iiesiiic, tiellscly

I' vcrrucose. Verrucac unilorrnly distributed, having a c1;irkish tinge, 2-3 each way, and probably 1i0110iv.

Unacet. 28 x 34 It, light brown witli a reddish tinge. Contents masked.

10. D. rubiginosa (Klf.) Iloorc (1503; N. America; Amcr. I.'ern SOC. no. 02). - Figs. 8, 13.

Spores 25 x 3 0 11; range, 20-28 x28-32 11. Ainb triangular with broadly rounded corners and straight to slightly convex sides. Prosimal face broadly conical, with protruding pole. Laesura arms 8 /L long. Esiric 2 It thick. Vcrrucac dense, up to 4 Iiigh, 5 It across, progressively larger to\var(is the distal pole, sometimes coalesced into irregular tubercles on the distal surface.

Unacct. 20 x25 11, light brown with a reddish tinge. Contents niaske(1.

11. 11. scabrci (Mrall.) IIoore (G77; T r i v a n d r u m , licrala, India; B. I<. N a y a r 4.1873). - Figs. 5, 15.

Spores 30 x42 p; range 24-32 x3G-18 11. Amb triangular with slightly concave sides. Prosiriial face nearly flat, with protruding pole; distal face convcs. Lacsura arms 16 /c long, their outer niargiris faintly thickened into smootli lips about 3 11 broad on either side of tlie laesura and tapering to\vards the tips. Esine light-brown, 2 /c thick, densely grnnulose and thickciicd here and there to form irregular vcrrucac (usually about 2 /L high and 4-8 /C

across, and having a reddish tinge) sonic of whicli coalcsce to form elongated ridges or even reticulations on the distal face.

Unacct. 30 x 38 11, nearly Iigalinc, containing a few sniall light-green plastids.

12. L). ienera (I'rcsl) Alctt. (1500; cult., EspLI. Gard.. Univ. I3russcls, I%clgium).

Spores 36 x 45 11; range 32-40 x 40-52 p. Prosinial face broadly conical. Laesura ariiis 18 /c long, with faintly tliickcned outer margins. Es ine deep brown, 2.5 ,I' thick. Scsine tliicker tlian nesine, densely and prominently granulosc and bearing sparse, irregular, vcrruca- to tuberculuni-like tliicken- ings (less tlian 2 p Iiigh), often irregularly coalesced and deeper in colour than the rcst of the esine. An outer skin-like layer of tlic csine, bearing the ornamentation (7 pcrinc) often cracks and peels off on acctolysis. Tlicrc is much variation in the ornanientation pattern, some spores having only a few csinous thickenings, whereas in others tlic thickenings are crowded, sometimes forming incomplete reticulate patterns.

Unacet. 28 x 38 11, nearly of the same colour as tlie acet., containing dense large light-green plastids. Some abnormal and ill-developed spores were noticed.

13. L). d / o r d i i Christ(OG1; cult., Bot. Gard., Univ. Tokyo). - Fig. 7.

Spores 40 x42 It; range 35-12 x36-48 11. Amb triangular with broad corners and nearly straight sides. 1-acsura arms 20 /c long. Esine light

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SI’OHE MOIII’IIOLOGT O F TIIE PTEIIIDACEAE. 111 191

4 7 8 = .

Figs. 1-8. Dicksonioitl and dennstacdtioid ferns (s, scsinc; n, nesinc). X I 1 figs. x 550, escept 1 I), . la, 511 and 8 b ( x 500) and 2 , 3 and 4 c ( x 1000). 1. Uicksonia librosa (a, prosirnal view, b, lateral vicw). - 2. D. sqttarrosa, sporoderni stratification. - 3. Cirlciia dttbin. optical section of a portion of the esine showing ornanientation. - 4. Cibolicm bnronielz (a, lateral view; b. prosinial view; c, sporodcrm stratification). - 5. Dennslaedlia scabra (a, prosinial view; I), latcral view). - G. L). appendicolafa. - 7 . 1). wil/ordii. - 8. I ) . robiginosa (a, proxinial view; b. lateral view).

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brown with a reddish tinge, 4.5 / c thick. Sesinc ii~ucli thicker tlian Iicxiilc, tlciiscly wrriicose. Vcrrucac irrcgular in outline, deep brown, 4 11 each (slightly sniallcr on the proximal facc) antl appearing liollow.

Unacet. 30 x36 11 and nearly of tlie same coloiir as the acet.

~ ~ I C I ~ O I , I < I ’ I . ~ I’rcsl

14. N. cciiiiliger~i Moore (445; Iinmaldiga, Ihmrup, Xssnm; 13. I<. Xagar & I’rcm Lnl 50979).

Spores trilcte, 25 x 37 11, wit11 little xx ia t ion in size. Amb triangular with broadly rounded coriicrs and sliglitly concave sides. Proximal face of spore nearly flat, with protruding angular pole; distal face convex. Lnesura arms 12 11 long, with tlic outer margins faintly thickened into sniootli lips about 3 11 broad oii either side of tlic Inesura and conspicuously tnpcring towards the tips.

I h i n e yellowish-brown, 1.5 11 thick. Sexine slightly tliickcr than ncxiiic, faintly and uniformly grnnulosc.

Unacet. nearly of tlic same size as acct., 1iy:ilinc to light-brownisli in colour, and with a few small, scattered, light-grccii plastids, oiie or two small ncarly liyaline oil globulcs antl Iiighly vacuolatcd protoplasm.

15. ,I[. linflongciisis Xngnr C? Iinur (1495; Hnfloiig, Assnm, India; 1’. Chnndrn 811,550). - Fig. 19.

Sporcs 26 x 34 11, with little variation in size. Ainb triangular with ncarly straiglit (somctiincs slightly concave or conres to prominently convcs) sides. Distal face of spore Iiemispherical to subconical. Laesnra arms 16 ji long. Scsinc nearly as thick as nesinc.

A fcw abnormal spores were noticed.

1G. ,lI. hcincei l’rantl (l5G9; Bircli Hill , Ilarjccling, 1iidi:i; 13. I<. Kngar 30717). - Figs. 16, ‘LO.

Spores 24 x 30 11; r:inge 22-26 x 36-14 11. Amb triangular with concave sitlcs :ind broadly roundcd corners. Prosimal face concave, distal face hcniisplicrical to subconical. Lncsura arms 18 11 long, their inner margins tliickcncd into proniinent smooth lips about 4 p broad on eitlicr side of tlic Iacsura and sharply tapering towards the tips. Es ine 2 11 thick.

17. df. Airla (Klf.) I’rcsl (1592; cult., Jnrd. ISsptl.. Unir. Brussels, I3elgiu 111).

Spores 3f(40-3) x 56 11; range 3 4 4 0 x 53-64 p. Arnb triangular with concave sidcs, bu t rarely with straight or convcs sitlcs. Prosimal face convex; distal face subconical. Lacsiira miis 18 11 long. Esinc very light brown, 3 p thick, denscly but wry faintly granulose.

Unacet. 32 x 40 i t , with tlic csine almost liyalinc.

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103

Figs. 9-18. Diclsonioid and dcnnstacdtioid fcrns. All x i50. 9. Dicksoniu anlarclicu. - 10. Culcila diibin. - 11. Cibofiurn glaucum. - 12. C . baromel:. - 13. Dennsfaedlia rribiginosn, distal view. - 14. I ) . appendiculofo, distal view. - 15. D . scnbra, distal view. - 1G. dIicrolepia Iinncei. - 17. Jl . plalyphylla. - 18. Lepfo- lepia novue-:elnndioe. 13 - 679514 Grnnn 1’111. vol. 8: 1

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18. ’\I. i,-icpeiiinsiiltie 11iirat:i (774; I<uzu\vata-slii, .\Iiii:iniata (:it\. Iiiimaiiiolo I’ref., Japan; 31. Kid0 3908).

Spores 31 x 44 p. Anib triangular with nearly straight to slightly coilc:lvc sidcs. I’rosirnal face concave, with the pole blunt or angular; distal face sublicmisplicricnl. Lacsiira arnis 16 / t long; lips about 4 / i broad on either side of the laesura. Esiiic liydinc to light brown with a darkish tinge, 3 thick, densely but very faintly granulosc.

and highly vacuolntcrl protoplasm.

* *

Unacct. 28 x 36 p. Spores containing inany small, light-green I)l:isti[ls

10. JI. kurzii (Clarke) I3cdtl. (101; Xgi~mbc, AIgsore, India ; 13. I<. Xnyar 4557G).

Spores 24 x 36 (10-1) 11; range 20-32 x 32-40 p. Anib triangular \\.it11 concave to nearly straight sides. I’rosimal face slightly concavc; distal face convcs to subconicill. Lacsura arms 16 / i long, their inner niargins tliickcnctl into faint, smooth lips tapering to tlic tips. Esinc light browi, 3.5 thick. Scsiiie iiearly as thick as nesinc, densely and prominently granulosc.

Unacct. nearly of the snnic size as tlic acct., bu t hyaline to pale-bro\vn in colour. Spores containing a few large greenish plastids. Scvcral ill-developed spores were noticed.

20. J I . nicirgiricila (Holtt,) C. Clir. (SOG; Iiuzuwata-slii, Minnmatn City, I i umamoto I’rcf.. Japnn; Al. Kitlo 3862).

Spores 40 x 58 p; rangc 36-42 x 52-6s I[ . Ainb tri:ingular with straight to slightly concave sides. Proximal face sliglitly concave; distal face sub- conical. Lacsura arms 1s / r long; lips 4 11 broad on citlicr side of tlic 1:icsura. Bsine almost Iiyalinc with :in yellowisli tinge, 3 thick, snlootli.

21. ill. plnti~pliyllu (Don) . I . Sin. (703; I)cVicolaiii, I(crala, India; 13. I i . Xayar 4529G). - Fig. 17.

Spores 25 x 42 1‘; rangc 24-2s x 36-4s I ( . Amb triangular (with straight t o slightly coricnre sides) to subcirculnr. Lacsura arms 16 16 Ioi~g. their inncr niargins faintly tliicliened. Esinc dccp-brown, 2 11 thick, smooth.

Unacet. 23 x 36 p, liyalinc with an ycllowisli tinge.

22. JI. p r o x i m i (Ill.) I’rcsl (1050; cult., Dot. Gards., Univ. Calif., 11 erlr eley, U. S .A).

Spores 40 x 50 11. Anib triangular with straight or slightly conyes sides. Distal fncc of spore licinisphcrical. Lacsura arms 22 ~t long; lips proniincnt, about 5 f t broad 011 either side of the lacsum. Es ine nearly hyaline, 3 /i thick. Scsiiie ncarly as thick as nexinc, smooth.

Unacet. contain many scattered, large, grccnisli plastids and a 1:irgc p:ilc-yellow oil globulc.

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24. Jf. S]JC[lfllCtlC (Linn.) lloorc var. iiillosissinici C. Clir. (1551; Jirkat:ing Camp, Soiith Xndaman Isls., G. S. Srivastavn 53140).

Sporcs 30 x 38 11; range 28-32 x 30-48 11. Ainb triangular wit11 strnigllt, slightly coiicavc or C O I I ~ C S sidcs (somctiiiics subcircular). I’rosiiiial face convex to ncnrlg Ilat; tlist:il lace srtblicmisplicricnl. Lncsura :irnis 13 1‘ long, tlicir iiiargiiis nnthickcnctl, 1:siiic tlecp l)ro\vn, 3 1‘ tliick; scsirie tliiiincr tli;iti

nesinc, densely grniiulose. Unacet. 28 x 34 11, light brosvii. AIaiiy :iborterl spores were noticed.

115. .If. s f r i p s “ (‘l’li~il~g,) I’rcsI (548; I’giiursl:i, I<. S .I. Hills, .4ssn111, 1mli:i; 1). li. X:iy:ir ii330).

Spores 30(3 1 -4) x .15(.19 -4) 11; range 2G-35 x 42-54 11. A i i b trinngular w i t h coiicavc sidcs. I’rosinial face concavc; distal facc so1)liciiiisl)licrical t o subconical. Lacsura ariiis 1G 11 long, their margins unthickciictl. Exilic light brow11 to nearly liy:iliiie, -1 11 tliiclr. The outer oriiaiiieiited layer of tlic esiiic sometimes cracks into picccs 011 :icctolysis.

Uiincct. 28 x3S p, hyaliiie, containing 2 or 3 large, nearly coloiirlcss to palc-ycllo\visli oil globulcs and n few sparse plastitls of various sizes. A fcw :ibortetl spores were tioticetl.

26- JI. forfrrgezisis Christ (Stis; Garo IIil ls , Xss:im, 1ndi:i; I$. I<.

Spores 2G x 3.1 11. Xmb tri:ingul:ir with str:iiglit or slightly coiivcs sidcs. Distal face of spore sitl~lic~iiisplicricnl to subconical. 1,acsura ariiis 12 long (sometimes sinuous), tlicir margins untliickcnctl. Esine hyaline with a brownish tinge, not clearly subdivided into scsiiie :ind iicsiiic, siiiootli.

Un:icet. nc:irly of the mine size atid colour as acet., contniniiig one or two large pale yellowish oil globulcs, a few sriiall scattered plastitls and liiglily vacuolatcd protoplasm.

i\‘ayar (i3433).

L 1 I\’ I) S AIS A 1)rgantlcr

25. L. eltiirzeii Ching (781; IIeluca, S:ita-AIara, Olisunii, Jnp:in; 11. Otomasu 48OG). - Fig. 25.

Sporcs trilctc, 18 x 36 11, with little variation i l l size. Arnb triangular with nearly straight sides and narrow corners. Prosimal face of spore nearly flat; distal face subliemispherical to subconical. Laesura arms 14 1 4 long,

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tlicir outcr margins tliickciictl into siiiootli lips about 4 11 broad (not tnpcrctl to tlic tips).

Exilic tlccp bro\vii, 3 ,I[ thick. Scsiiic tliickcr thnii iicsinc, tlcnscly riigulose with the rugulne up to 2 11 high, 2 11 bro:itl, 3-5 11 lorig and irrcgulnr in slinpc.

28. L. cirllrrrlrr (\\'illd.) Sn-. ( 5 i i ; Sliillong l'cal;, .4ss:ini, Iiidia; 13. Ii. Xnynr tij150-l). - Fig. 21.

Spores iiionolctc, bilateral 32 x 48 x 30 11, with little vnrintioii in sizc, plnnocoiiws (lnt.), oblong (pol.). Lncsirrn 22 11 long, its outcr innrgiiis proiiiiiiciitly tliickeiicd into siiiootli iiniforrii lips about 3 11 brontl oil eithcr side of the Incsura and protrutliiig G-8 i i from thc sclcriiie surface. Sclcrinc brown, 2.5 11 thick, faintly grniiulosc. A thin, outcr, skin-like laycr of tlic sclcriiic (? pcrinc) oftcii cracks irrcgulnrly into siiinll picccs nntl soiiietiiiies peels off on ncetolysis.

Uiiacct. iicnrly of tlie sniiic size ant1 colour ns tlic ncct., containing innny lnrgc crowtlcd plnstids.

19. I,. cleconi~miltr l\'illtl. (1 SO; I'yiiiirsla, 1;. c!! .J . Hills, Ass:iiii, India; 11. I<. Xnyar 49813).

Spores moiiolctc, bilnternl, 37 x 50 x 38 11; range 32-40 x 5G-72 x 3G-40 11,

plniiocoiivcs (Int.), oblong (pol.). L:icsrirn 28 11 long, its outcr iiinrgiiis tliickcnctl into proininent, protrutlcd, smooth lips about 3 11 broad on citlicr sitlc of tlic Incsurn. Esiiie light browii, I 11 thick. Scsiric iiiucli thicker than nesinc, faintly but tlcriscly riigiilosc (rugulne lcss tlinii 2 11 tall, 3 11 broad, 4-8 11 loiig) with thc rugulnc soriictinics conlcscctl.

Unacct. 28 x 40 x 28 11 in sizc, iicnrly Iiyalinc with nn ycllo\visli tinge, mid contnining riiany Inrgc, dense plastitls.

30. I,. lirieriris S\v. (1 555; Gordon, Xc\v South \\'ales, Awstralia; Knspicw 2-1-99).

Sporcs 28 x 4.1 ,I(; rnngc 24-30 x 40-48 11. Aiiib triniigulnr \villi stmiglit sidcs :ind broadly rouiidetl coriicrs. Lnesura :irnis 1 G /i long; lips about G / i

broad on eitlier side of tlic lncsura and conspicuously tnpcriiig towards tlie tips. Sclcrinc brown. Scsine slightly tliickcr tlinn ncsinc, dcnscly and proiiii- ncntly graiiulosc. An outcr skin-like layer of tlic sclcriiic (? pcrinc) 1)c:iring the graiiulnc oftcii cracks and pcels off on ncetolysis, Icaving n siiiootli sii r f ace.

Uiiacct. ncnrly of tlie sanie sizc, b u t lighter brown (with an ycllo\visli tinge), ant1 containing two or tlircc smnll, nearly hynlinc, oil glol~ulcs nntl rnniiy Inrgc, crowded plnstitls.

31. I.. lobnla Poir. (G2-l; Gnro IIills. Assnm. India; 13. I i . Snynr G34-15). - Fig. 22.

Sporcs 23 x30 p, with little variation in size. Lncsura arnis 12 11 long; lips about 2 ,IL broad on eitlier side of tlic Inesorn. Esine light brown with n

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blackish huc. Scsinc iicarly as thick as iiesine, densely and prominelltly granulose.

Unacet. tiearly of the s:imc sizc and coloiir as tlic acet., containing lilally large crowded plastids.

32. I,. n~irrophylla Su-. (1544; School biol. Sci., Univ. Syd~icy). - Fig. 26.

Spores 28 x 40 j i ; range 24-36 x 36-44 11. Amb triangular witli straiglit or slightly convex sides. Prosimal facc nearly f la t t o slightly convex; distal face hcniisphcrical. Lacsiira arms 10 ji long. Sclerinc brown, 3.5 j i thick, dcnscly aiid promincntly granulosc, An outcr, skin-like, ornamcntcd layer of the sclcriiic (? pcrine) cracks and sometimes peels off, leaving a smootll surfnce.

S I’H EX 0 ;\I EIII S ;\I nson

33. S . billom (]<If.) Ching (796; I\lanazurii-Cape, I<anaga\va I ’ d . , Japan; I<. Lidn s.11.).

Spores nionoletc, bilateral, 30 x 48 x 36 p, with littlc variation in sizc, p1:inoconvcs (Iat.) oblong (pol.). Lacsura 36 /i long, its outer lllargiiis thickened into prominent sinootli protruded lips about 2 p broad on eitlicr side of tlie laesura.

Esilic deep brown, 4 11 thick. Sesine thinner tlian ncsine, sniootli. Pcrine (kirk brown, not clcarly tliffcrcntiated, tlenscly grnnulosc, closcly atllicrcnt to csinc and skin-like hut oftcri cracking and pcclitig off in pieces.

3-1. S. clruscrm (Linii.) Copcl. (395; ?‘urn, Gnro Hills, Xssnni, India; 1%. I;. Xayar 50919). - Fig. 23.

Spores 28 x 4 5 x20 j i , with littlc variation in size. Laesura 22 j i long; lips about 1.5 11 broad on either side and tapering to either end. Exilic 3 ji thick, not clearly diffcrcntiaterl into sesinc and ncsinc. Pcrinc skin-like partially pccling off 011 acetolysis.

Unacct. ncarly of t he samc sizc as the acct., bu t lighter in colour, atid contaiiiiiig inany large plastids and a large ycllowisli oil globule.

SC I I I %O I,O A I X Gait ( I . 35. S . ensi[oliim (S\v.) J . Sin. (1 129; Castle IIocl;, Algsorc. India;

1’. Chantlrn 95129). - Fig. 28. Sporcs trilctc, 32 x 3 9 1‘; raiigc 26-36 ~ 3 2 4 . 1 p. Amb triangular with

nearly straiglit sides and broadly rounded corners. Prosimal facc of sporc broadly conical; distal facc subhemispherical to subconic:il. Laesura a r m 12 11 long, tlicir outcr margins thickened into smootli, uiiiforin lips about 2 j i broad on citlicr side of tlie Iaesura, but not vcry distinct towards tlic tips.

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S PO I% E JI 0 1% 1’1 10 1.0 CY 0 I: TI I I: I’T I: 111 1) .\C I::\ I:. I I I 199

Esinc brown, 4 11 thick. Scsine nearly as thick as liesine, sniootli. I’erine not clearly differentiated, skin-like, closely adherent to the csinc, cracking and peeling off in picccs on acctolgsis, and tlcnscly rugulosc. Ilugulac irregular in outlinc, 1 11 high, 1 11 broad, and about 2.5 r i long.

Unacct. 25 x 3-1 /i in size, lighter in colour than the acet., and containing a fcw, scattered, large, light-green plastids.

3G. S. hclcrophyllnni (Drynndcr) J. Sm. (1 ‘718; Cnstlc Itocli, Mysore, India; 1’. Chniitlrn 01900).

Spores 32 x 40 11. Ainb triangular with slightly conves sides and rather iiarrow corncrs. Prosinial face of spore nearly flat. Lncsura arms 20 11 long. Esinc tlcep brown, 3 11 thick. Perinc densely granulose.

TX 1.; X I T I S \\‘i I 1 (1.

- Fig. 30. 3’7. T. sp. (1372; Clicrrnpunjcc, Xssam, India; 1’. Cliniitlrn 8001 1) .

Sporcs trilctc 28 x 37 11; range 24-32 x 32-44 11. Amb triangular (with coiivcs sides) to subcircular. Proximal face of sporc convcs, dist:il face subliernisplierical. Lacsura arms 1s / t long, thcir outer margins tliickcncd into promincnt, smooth lips about 3 / L broad 011 either side of t he laesura and protruding conspicuously from the general surface of the esinc.

Esinc dccp brown, 2 /L thick, not clearly diffcrcntiatctl into scsine and nesine, tlcnsely verrucose. \’crruc:w subglobosc 2-3 11 across, 2 high.

Alany abnormal sporcs, soiiic ill-dcvclopcd and crumpled and others csccptioiinlly Iargc, w r c obscrvctl.

LEI”I’OLE1’IA JIctt.

38. I,. nooccc-:clrrntlirie (Col.) I<ulin (1 588; Xc\v Zcalnnd; Xiiicr. Fern Soc. 110. 115). - Fig. 18.

Spores trilete 22 x 32 11; rangc 20-26 x 2s-36 11. Amb triangular with ncarly straight to slightly coiicavc sidcs and broadly rounded corners. I’rosimal face of sporc concave to ne:irly flat; distal facc srrblicniisp1ieric:iI to subconicnl. Laesura arms 12 11 long, tlicir margins untliickcnetl.

Esine pnlc-brown, 2 /L thick, not clearly diffcrcntiated into scsine and ncsinc, smooth.

Unacct. 20 x 26 p, liyaline to light brownish, containing inany small light-green plastids and a few small yellowish oil globulcs. Alany abnormal and iindeveloped sporcs wcre obscrvctl.

I ~ Y I ~ O I X P I S Bcrnli.

J1ntlr:is. India; 13. I<. Nayar 45384). - Figs. 24, 29. 39. If. p i m c f n f a (Thnbg.) Alett. ( 1 27; I’alparni, Coiml)atorc,

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Spores nioiiolctc, bilntcral, 25 x 37 x 2s 1 4 ; rangc 20-2s x 32-40 x 24-30 planoconvcs (ht.), ol)long tciitling to he subcircular (pol.). Lncsura i j loiig, its iiiiicr iiiargiiis faintly tliickciicd.

iicsiiic, spiiiosc. Spincs about 5 / i lbiig, 3 / r thick, sonictinics fl:ittciicd (6 broad).

Uiiacct. 2.1 x 32 x 20 11, iicarly liyaliiic in colour, containing a fc\v ligllt- green plastids, one or two nearly hgnliiie oil globules and conspicuously v:icuolatctl pro toplnsni.

. Esine light ycllo\visli-bro\vn, 3.5 11 thick. Scsiiie ncnrly :is thick :Is

Sporcs 20 x 2G x 20 11, with little variation in size, clongatc-oblong (poi.). Lacsum 1 G 1‘ loiig, its margins uiitliickeiied. Esiiie gcllo\\-isli brown, 2.5 thick, dcnscly spinosc. Spincs 2 / i thick, up to 5 / c high, either blunt or rather sharp, sometimes more or less f1:ittcnctl (up to 4 / i bro:id) aiitl oc- casionnlly ciirvcd towards the tip.

4 1. H. sprsisorcr (Scliratl.) Iiuhii (12%; I’ictoria, E. Xigcrin, 1’. \\’. Itichnrtls 3G52G; FHI. ) .

Spores 20 x 3-1 x 20 11; raiigc 16-24 x 24-36 x 1G-24 p. Exilic diiIl-bro\y11, 3 p thick. Scsinc inucli thicker tliaii ncsiiic, dcnscly spinulosc. Spinulcs ninniinillatc, 2 14 tall, and soiiictiincs rntlicr flattciictl aiitl rugulalikc.

Unacct. nearly of tlic same size as tlic acct., aliiiost hgaliiic, coiitaiiiing a few sparse plastids.

I’TP~RI DIUM Scopoli -42. Z’. (rquilinuni (Linn.) Iiulin ( G i ; ~lauplilang, I<. & J. Hills,

ASS~III ; India; 13. Ii. Xaynr & I”?m La1 50476). Spores trilctc, 25 x 37 I t ; rangc 20-27 x 32-40 11. Amb triangular with

rather coiivcs sides. Prosimal face of spore flat; distal face convcs. Lnesurn arins 16 /i long, their iiiargiiis not tliickciicd.

Es inc deep-brown, 2.5 11 thick. Scsinc thickcr than nesinc, tlciiscly mid promincntly grnnulosc. Grnnulcs Iicmisphcrical, about 1.5 1‘ cacli way mid tlarkcr in colour tliaii the rest of tlic csinc. A thin outcr layer of the csinc rarely cracks aiid pcels off in pieces.

Unncct. 22 x32 p, nearly of the sainc colour as tlic ncct. Contcnts iiiaskcd.

43. 1’. r r q i d i r i t r n i ( L i i i i i . ) I<uhn var. Iriniyinosuni Cliilc. ( 1 370; I3cl- ton mo., Missouri; L. 31. Uiribacli ‘732).

Spores 22 x 3 5 ,I(; rnngc 20-24 x28-14 11. Amb triangular with iiearlg straight sides and iiarrow coriicrs. Lacsura arms 10 long, with the outcr margins thickencd into narrow (about 1 broad on eitlicr side of the Incsurn) uniform lips. Esinc dccp brown, 1.5 /c thick, not c1e:irly dilfcrcntintcd into

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20 1

Figs. 2G-31. 1,intlsrryoirl and Iiypolcpitloitl ferns. 2\11 figs. x 750. 2G. Lintisticcc microphglla. - 27. L . chiineii. - 28. Sciiizoloma erisi/oiiiim. - 29. I I r~po- lepis piiiicfala, polar view. - 30. Toenif is sp., distal vicw. - 31. Ilisliopleris iiicisn, lateral view.

scsiiic arid ncsinc, grnnulose. Grnnulcs somctimcs coalescing to foriii short rugulnc.

Unncct. nearly of tlic sniric sizc as the acct., light-hrown in colour ant1 contnining n lnrgc, nlniost Iiynliiic oil globulc.

1 I I S‘I’IO PTER I S J . Sin.

4-1. H . incisrt (‘Hiiibg.) J. Sin. ( 3 3 i ; Darugiri, Gnro Hills, Assam, Ind in; 11. I<. X:tynr G3.137). Figs. 25, 31.

Spores nionolctc, bilntcral, 30 x 48 x 3G 11; range 2s-3G x 44-56 x 24-32 p, planoconvcs (Int.), clongntc-ovntc (pol.). Lacsura 23 /i long, its outcr margins faintly thickened into lips about 2.5 11 bro:id on citlicr sidc of the lncsurn mid with undulnting surfncc.

Esine dull-brown, 3 / i thick, not clearly tliffcrcntintcd into scsiric and ncsinc, tlcriscly tubcrculntc. Tubercles crowlctl, sonietiincs conlcsccd to form rugulnc with undulating surfncc and sides, larger (about 4 ,I‘ high, 10 11 across) and iiiorc promincnt on thc distal surface of thc spore but pro- gressively less prominent towards the prosirnnl polc and almost nbscnt nroiiiitl the lncsura.

Unncct. 23 x -10 x 23 11, almost ilynlinc to liglit ycllowisli-l)rowii in colour, and containing many sm:ill light-grccn plnstids and proniincntly vacuolntctl protoplasm. AInny :ibnornial spores occur (sonic of tlicni abortctl and cruniplctl, ant1 otlicrs csccssiwly Inrgc).

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Sutiiniaty

Spores of 44 spccics (14 gcncra) 1)clonging to the dicksonioitl, dcnnstncdtioitl ant1 the lintlsnyoitl groups of tlic I’tcritlaccae ( s e ~ l s i ~ Copcland) arc dcscribcd from acctolyzcd arid frcsli samplcs. JIono- lctc, hilatcral spores occur in Hisliopteris, Hypolepis ant1 Sphe~~orrieri .~. In tlic othcr genera thc spores are of the trilctc tctrahcdral type, csccpt in Linrlscieci in which hot11 types occur. X \vcll-tliffcrcntint~~l pcrinc sccins to hc absent, except in the lindsayoid ferns in hi& a thin, sliin-like, ill-tliffcrentiatcd laycr of tlic sporc-coat (? perinc) often cracks nntl soriiclinics pccls on in some species. Aniong thc tlicksonioitl ferns thc csiiic is areolatc in Diclisoriin ant1 Ciilcitci, smooth in Orthiopteris and tubcrculntc in Ciboliiini. In the Dcnn- stacdtioidcnc, Jlicrolepici ~)osscsscs a grnnulosc csinc with 1:irgc grnnulcs (ca. 1.5 p across) in some spccics, \vliilc Dcririslrrcdfici is charnctcrisctl by n verrucosc csinc. ?‘lie sclcrine is commonly grnnulosc or sonictimcs rugulosc in the lindsayoitl ferns, except in Trcenilis in \vliicli tlic cxcrcscenccs arc rather large (en. 2 14 across) ant1 vcrroca-lilic. I n Hypolepis and Pleritliiim tlie csiric is spinulosc to spinosc lvhcrcns in Ifisliopleris it is tul)crculatc. A distinct cqiia- torinl collar-like ridge occurs in Ciboliiini. Fresh spores in all gcncra contain many pnlc-grccn plaslitls. I n most cases the spores swcll on acctolysis. ‘The csinc of Dicksonirr sqiicirrosci dissolves in the acctolysis mixture.

Tlic spores of the dic1;sonioid ferns arc niarlictlly larger tl inn thosc of thc otlicrs (40-4G x 50-70 p). Orlliiopleris is exceptional in the group, possessing sniallcr spores similar to thosc of tlic dcnn- stnedtioitls in which the avcragc sizc wr ies from 25-40 x 30-58 p. Comparatiwly siiinllcr spores clinrnctcrisc the lindsnyoitl ferns, Hisliopicris, IIgpolcpis, anti Pteridiiim.

Acknowledgements

\Ire arc tliankful to Jlr. 1’. C. Iloy and Alr. JI. Ii . Tantlon for their help in thc prcpnrntion of tlic illustrations and to the various agcncics mcntionctl in the tcst for supplying sonic of tlic spore s:imples. I t is a plcasiirc to rccortl liere our gratitude to the. Director, National Ihtnnic Gartlcns, for his cncourngement.

Nation nl 13 o t n ni c G art1 c ns, Luck n o\v

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RE FE I t ENC E S

A u r ~ ~ i i ~ i r , A., Krs~s, C. A., and AlATiiEw, 1’. AI. 1962. Stutlics on tlic cyto- logy and pliglogcny of the Pteridopliytes. 1’11. Observations on one hundred spccics of South Indian Ferns. - J. Indian bot. Soe. 11:

Ciilsc;, 11. C. 1940. On natural classification of I’olypotliaccae. - Sunynt-

CII~ISTESSES, C. 1936. Filieincae. - Chapter SS, in \‘erdoorn’s “I\Ianual

COPELASD, E. B. 19-17. Gcnera Filicnm. - \1’altliain, Akiss., U.S.A. ~ I O L T T U ~ I , R. E. 19-17. A reviscd classification of the leptosporanginte fcrns. - J. Linn. SOC. (Bot.) 53: 123-158.

MASTOS, 1. 1958. Cliromosonics and fern pliylogeny with special reference to “I’tcridaceae”. - J. Linn. SOC. (13ot.) 11: ‘i3-92.

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scnin 5: 201-2GS.

of Ptcridology”. - Tlie Hague.

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