Palynological study in Aspleniaceae from North-West Argentina - Sociedad Argentina de Botánica

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Bol.Soc. Argent.Bot. 37 (3-4): 217

-

229.2002

PALYNOLOGICAL

STUDYIN

ASPLENIACEAE

FROM

NORTH-WEST

ARGENTINA

G.E. GIUDICE1,M. A.MORBELLI2AND M. R.PINEIRO2

Summary: Thespores of Asplenium L. species fromNorth-West Argentinawereanalysedunderlight

and scanning electronmicroscopes. Thesporesare monolete and 40-50pm Indiameter;the exospore

Is 1.2-2.5 pm thick,one-layered insection and smooth. Theperisporeis 1 to 10pm thick,seeminglyone

layered with three-strata insection and çamerate. The elements ofthe sculptureareechinulae,crlstae,

rugulaeandridges. Correlationbetween spore size andploidylevelswas found, sincespecies referred

as tetraploids andhexaploids produce largerspores.Onthe basisofthe combination of characters of the outer perispore stratum, suchasfolds(typeanddistribution), degreeof perforation,andsculpture

(type,locationandcombination),the species presentintheareaofstudywerecategorisedintotwo maingroups: wingedand ridged.Speciescan be determinedbytheirspore ornamentation. Most species

fallInto thewinged, scarcely perforatetype, with varied ornamentation between folds. Nocongruence

was foundbetween spore characteristics andvegetative characteristics, with theexception of

Aspleniumserra,inwhich both characteristics makeits determination possible. Key words:Asplenium,spores,perispore, exospore,Argentina.

Resumen: Estudio palinológicoen las Aspleniaceae del Noroeste deArgentina. Seanalizaroncon microscopio ópticoy electrónico de barrido lasesporas de las especies de AspleniumL.quecrecen en

elNoroestedeArgentina.Lasesporassonmonoletes, de 40-50 pm de diámetro, conexosporiode 1.2-2.5 pmde espesor,deuna capa en sección y liso.Elperisporio, de 1a 10 pm de espesor, posee

aparentementeunacapacon tres estratos en secciónyes camerado. Loselementosde la escultura son espínulas, crestas,rúgulasylomos. Seha observado correlación entre eltamañode lasesporas y

el nivelde ploidía, ya que especies citadascomo tetraploides y hexaploides poseen lasesporas de mayor tamaño. Enbasea la combinación de caracteresdel estratoexternodel perisporio, talescomo:

pliegues(tipo y distribución),grado de perforacióny escultura(tipo, ubicación ycombinación), las

especiespresentesenel área deestudio pueden ser reunidas en dosgrupos:aladas y lomadas. La

mayoríade las especiesse encuentrandentro deltipoaladocon escasasperforacionesyornamenta¬ ciónvariadaubicada entre pliegues.No se ha observado congruencia entrelos caracteres de las

esporas ylos vegetativos, con excepción deAspleniumserra, en laque la combinación de ambos caracteres permitesudeterminación.

Palabras clave: Asplenium,esporas, perisporio,exosporio, Argentina.

Species of sections Asplenium and

Sphenopieris

fromNorth-West Argentina, have free venation and aredifferentiated by their bladestexture, sorusloca¬ tion and rachis indument (Morton & Lellinger,1966; Sota, 1977).The sporecharacteristicswerenot men¬

tioned in these studies.

AccordingtoMurakami&Moran(1993), Asple¬

niumlaetum Sw. (sectionHymenasplenium) is widely distributed and theonly representated of that sec¬ tion which reaches theNorth-West ofArgentina. The

authors characterised the spore of section

Hymenaspleniumasechinate,papiloseorcristateas inA. laetum.

Thesporemorphology oíAspleniumwasquoted in floristiccontributions like those of Tryon & Tryon (1982)and Murakami&Moran (1993)and, in studies where morphology andultrastructurewere studied indetail,suchusthose ofNayar & Devi (1964) and

INTRODUCTION

Thesporesof the Aspleniaceaeareanalysedas

part of the project Palynological Flora of the

Pteridophyta ofNorth- WestArgentina. This family is representedby Asplenium L.:A. achólenseHieron.,

A. argentinum Hieron., A. auritum Sw., A.

depauperatum Fée, A.

_formosum

Willd.,A. gilliesii Hook, A. laetum Sw., A. lilloanum de la Sota, A. lorentziiHieron.,A. monanthes L., A.palmeri Willd., A.praemorsum Sw., A.pumilum Sw., A. resiliens Kunze, A.serraLangsd., A.triphyllumPresl and A.

tucumanenseHieron. (Sota, 1977; Ponce, 1996).

Cátedrade MorfologíaVegetale-mail: [email protected]

2Cátedra

de Palinologiae-mail: [email protected], Facultad de CienciasNaturalesy Museo, Universidad Na¬ cional de La Plata, Paseodel Bosque s/n°,1900,La Plata, Argentina.

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(1960) aftertreatmentwithhot 3% sodium carbonate for 2 minutes. For SEM, the materialwastreated with hot 3% sodiumcarbonate, washed,dehydrated,sus¬

pended in 96% ethanol andthen transferredto

acetateplatesand coated withgold.Inordertostudy the ultrastructureusingthe SEM wall fractureswere made withultrasound.

The observationsweremadewithaJEOL

JSMT-100 scanning electron microscopeat Museo de

CienciasNaturalesdeLaPlata.

The sporesmeasures wereestimatedon25 spores ineachsamplequoted inMaterial and Methods, and given inpm.The figures correspondtominimum, average (in brackets) and maximum values respec¬ tively. The values of the maindiametersinclude the perispore thickness.

Theterms andconceptsproposedby Viane & VanCotthem (1977)

,

Puttock &

Quinn

(1980)and Tryon &Lugardon(1991)wereusedtodescribe the perispore features and structure.

Nayaretal. (1964), Viane & VanCotthem(1977), Puttock &

Quinn

(1980), Perez Rayaetal. (1986), Pangua & Prada (1988),Prada etal. (1989), Braggins & Large (1990), Tryon (1990), Tryon & Lugardon (1991)and Johns (2000).

Withrespecttothe species that growinthearea

of study,A.formosum,A. praemorsum, A.auritum, A. monanthes and A.serrawereillustrated and de¬ scribedby Prada etal. (1989) basedonmaterial from

Colombia andPerú,those ofA. monanthes, A.

resiliens,A.serraand A.auritumwere illustrated with SEMphotographsin Tryon & Tryon (1982) and Tryon &Lugardon(1991) and, those of A. laetum and A.pumilumin Tryon &Lugardon(1991).

Michelena (1993) studied the speciesof

Asple-niumthat grow inBuenos Aires Province and

Morbelli(1980)studied those ofPatagonia. Inthe worksofPuttock&

Quinn

(1980)andBraggins &Large (1990 ), theperisporefeatureswereanalysed and taken intoaccountinordertoestablish the spore typesinspecies ofAustraliaandNew Zealand.

Tryon(1990) founda correlation between the sporeornamentationand the ecologyinspecies of Asplenium. She found thatepiphytic species have morecomplexity in the spore sculpture than those of terrestrial andepilithic species.

In Loveetal. (1977) and Tryon & Tryon (1982) referencestocytologicallevelsinAspleniumwere

found.

It wasobserved that polyploidy is frequent in speciesthatgrowin AmericaaswellasinEurope. This

characteristic

wasrelatedwithanincrease in .sporesize (Perez Rayaetal., 1986;Tryon & Tryon,

/.c.; Tryon, 1990).

Ouraim in this study istoanalysethe spores of the seventeenspeciespresentinNorth -WestArgentina

under lightmicroscopy and scanningelectron

microscopyinordertoprovidemoreinformationabout the species that formpartofFloraprojects inour coun¬ try.With thatinformationwewilltrytofindoutthe characteristics sharedby allthe speciesaswellasthose that could beusefulforsystematicpurposes.

Specimens Investigated

The letters MP in this list and the figure legends indicate the reference number of each pafynological sample which is filedat in the Laboratorio de Palinología,Facultad de CienciasNaturales and MuseodeLaPlata.

Asplenium achólense

Prov. Tucumán: Dpto. Monteros,

Qda.

Pueblo

Viejo, De la Sota 4060, 1-1965(LP), MP 3733.Prov. Salta:Dpto. La Caldera, AltoLaSierra, camino de comisa,alt. 1400m.,Palací 1036(MCNS),MP 3612

.

Asplenium argentinum

Prov. Jujuy:Dpto.Capital,C°Labrado,De laSota 4368, 11-III-1966 (LP);MP3741.;M>w, Yala.

Sotelo s/n, 9-VI-1948 (LP); MP 3751., Dpto. Ledesma, caminoaValle Grande;Cabrera & Fabris 22620, 13-V-1972 (LP); MP3752. Prov. Salta:Dpto. Capital, Loe.

Qda.

San Lorenzo, alt. 1600 m.s.m., (

MCNS 167);MP 3611.;Dpto.Capital,Loe.

Qda.

de

SanLorenzo, alt. 1600m.s.m.,MCNS,Nro.168; MP 3610.;Sta. Victoria, 1400m.s.m.,Mármoletal.,9205c, 21-IX-1972; MP3743.;Dpto. Sta. Victoria, caminoa The workwasbasedonherbarium material from Baritú,1530m.s.m.Marmol,Cuezzo & Cuezzo (h),

18-the

followinginstitutions: BA, CT, LP, LIL and SI. IX-1972 (LP); MP 3754; Prov. Tucumán: Dpto.

The sporeswerestudiedusinglight microscope (LM) Monteros,

Qda.

Pueblo Viejo,Dela Sota 4075,1-1965 andscanningelectron microscopy(_SEM). For LM, (LP); MP3742; Dpto.

_Tafi

delValle,Yerba Buena, the materialwasacetolyzedaccordingtoErdtman Venturi184,15-1919 (LP);MP3753.

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Asplenium auritum Asplenium monanthes

Jujuy:Dpto. ValleGrande, serranía de Calilegua, Prov. Jujuy: Dpto. Capital, El Cucho, De la Sota

4311, 9-III-1966 (LP);MP3744;Dpto.Ledesma,Valle Fabris,Crisci&Paridla5933,15-10/II/1965 (LP);MP Grande, Cabrera &Kiesling 20319, 11-XII-1969 (LP). 3746; Dpto. Capital, Lagunas de Yala, Cabrera Prov.Salta:Dpto. Orón, Loe. AguasBlancas,fincaEl 16365, 28-X-1964 (LP).Salta:Dpto. Capital,

Qda.

de

Arrazayal,alt. 850m,MCNSNro. 91;MP3613;Dpto. SanLorenzo, alt. 1600m.s.m.,MCNS,Nro. 158; MP

LaCaldera,Loe. camino decomisa,AltoLaSierra, 3624; Dpto. Rosario deLerma, Loe. 2km pasando 1400m,MCNSNro.1037;MP3614;/¿/ew,Loe.Ruta9, Corralito, caminoaEl Manzano, alt. 1750m.s.m, camino de comisaaJujuy, 8kmarriba de Santa Laura, MCNS, Nro. 3537, MP 3623. Tucumán:La

Cueva,

alt. 1500m.s.m., Novara,MCNSNro. 3276;MP3615.; Rodríguez483, 3-IV-1912(LP);MP 3747.

Prov. Tucumán: Dpto.

_Tafi

del Valle,Yerba Buena, _{Asplenium palmeri} Venturi201

,

21-II-1919(LP);MP3745.;Dpto. Monteros,

Eda.PuebloViejo,Dela Sota4073,1-1965 (LP) _{Brown 525 (LP),MP 3860.}Salta:Dpto.Anta,P.N. ElRey, Ayo. LosPuestos, Asplenium depauperatum

Prov. Catamarca:Dpto. Belén,El Rodeo, Asplenium_Jujuy:Dpto.praemorsum_{Capital,ElCucho, De la} Sota4335,10-Castillón2018, 15-1-1911 (LIL); MP3765;FW. Salta:

_m-

_{1966 (LP);}_{MP 3748.}_{Salta:Dpto.La Caldera, Loe.} Dpto.Capital, Loe. Cerro San José (Río Wiema), alt. _{Ruta g} camjno_{de comisa}_a_{Jujuy, 8 km}_antes_{de Abra}

1300 m.s.m.,MCNS 604;MP 3616;Prov. Salta:Dpto. _Santa

_Laura?

_{1500 m.s.m.; Nro.}_3278; _MP_3622;

Rosariode la Frontera, Candelaria, Venturi3837,

14-IV-1925 (LIL);MP3767. Dpto._{19-1-1979 (LP);MP 3750;Dpto. Orán, Aguas}Anta,“ElRey”,caminoaPozoVerde,Brown437,_Blancas,

Palací 102, 24-VII-85 (LP); MP 3755. Tucumán:

Prov. Salta: Dpto. Orán, RíoPescado,.Abra Dpto.

_Tafi

delValle,Venturi 277,III-1919(LP);MP3749. Arasayal, Vervoorst & Cuezzo 7599c, 26-X-1970; MP

3762;Idem,PuestoRivero,Hiuck 127, V-1950

(LlL),

MP3769;idem,RíoPescado,Vervoorst&Cuezzo 7808c,

25-X-1970(LIL),MP3770.Prov.Tucumán:

_Qiíebrada

_A¡taÿo_{Salí, Schreiter}_8,_{2-V-1923 (LIL);MP3768.} de Lules,Castillón 11668,Vin-1911(LIL);MP3761.

Asplenium

_formosum

Aspleniumpumilum

Tucumán:Dpto.Capital,BarrancaColorada,Venturi 817(LP), 5-V-1920; MP 3734

_-

MP3760;Dpto. Cruz

Asplenium resiliens

Salta:Dpto. Anta,P.N.ElRey,Brown985(1) (LP), 19-7-1979;MP3735.Jujuy:Dpto. Valle Grande, Valle Agarde 477,21-1-95 (LIL),MP3766.Prov.Salta:Dpto. _Colorado,_Kiesling_et_{al. 541 (LP), 21-1-1974; MP} Cachi,Loe.ValleEncantado,alt. 3000m.s.m.,MCNS _3736._{Dpto Capital,}_El_{Cucho, C°}_{Labrado, De}

_1ÿ

Nro. 966; MP 3617. Prov. Tucumán: Dpto.

_Tafi

del _Sota4379_aP\ _\_\_nI__{1966;MP 3737}

Valle,Castillón 644, 26-XII-1907 (LIL),MP3764. Asplenium gilliesii

Prov. Jujuy:Dpto. Valle Grande, C° Hermoso,

Aspleniumserra

Asplenium laetum _Jujuy:_Dpto._Ledesma,_camino_a_{Abra de Cañas,}

Prov.Salta:Dpto. Orán,Río Pescado,AbraArasayal, ]_000.1700_m.s.m.,_De_{la Sota 4410 (LP), 17-III-1966;} Vervoorst& Cuezzo7598c,26-X-l970(LIL),MP3763. _MP₃₇₃₈_{_Salta:}_Dpto._{Orán, Aguas}_Blancas,₁₂₀₀ m.s.m.,Palací 111 (LP), 24-VIII-85; MP 3739;Dpto. Prov. Salta: Dpto. Anta,P.N.E1Rey, Brown865 SantaVictoria, camino de BaritúaPorengal,Cuezzo, (1),9-VII-1979;MP 3783

Aspleniumlilloanum

Cuezzo(h)&Mármol9277c(LP),18-IX-1972;MP3740. Asplenium triphyllum

Jujuy:Dpto.Rinconada,Mina Pirquitas,l-III-Aspleniumlorentzii

Prov. Catamarca: Dpto. Belén, Potrero Ambito,

Qda.

delNacimiento,LIL 41534;MP3621

.

Prov.Jujuy: 1964, Schwabe 905 (LP);MP 458 Dpto.Capital,Loe. Lagunas deYala,LIL 187938, 10- Aspleniumtucumanense

01-47; MP 3620. Salta:Dpto. Capital, Loe.

Qda.

de Jujuy:Dpto. Capital, S.S de Jujuy, Parque

SanLorenzo,alt. 1500m.s.m.,MCNSNro. 130, 11-03- Botánico,Vázquez 82 (Fac.Cs. Agrarias),15-JV-1987;

94; MP 3626;

Qda.

de San Lorenzo, alt. 1500-2000 MP3585; Dpto. Ledesma, caminoaValle Grande,

m.s.m.,MCNSNro. 165,25-08-85; MP3627;Dpto. Abra de Cañas, Cabreraetal. 25679 (LP), 31-X- 1974; RosariodeLerma, Campo Quijano, alt. 1550m.s.m., MP3581;Dpto. ValleGrande, Vervoorst & Cuezzo

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The perispore generallyappears as one-layered Brown102(1) (LP), 18-VII-1979; MP3584; Dpto.

Chuquisaca, Monte Grande, Saravia Toledo 11909 with three strata insection and is camerate. The (CTES),20-VI-1993; MP3583; Tucumán:Dpto. innerstratum(Pi) isthin andattachedtotheexospore. Famailláyilla Nougués,Venturi s/n° (GH), 23- V- The middlestratum(Pm) is basicallyacontinuous 1922; MP3669;Bajo de Anfama,Lillo 5036 (GH), 8- spacefdled with radialpillars (rodlets) thatcanalso

VI-1906;MP3668. besingle, digitateorconcrescent.Thepillarsare more

evident atLMinAsplenium achalense.Theouter

stratum(Po)is thin and generally perforate; it bears the elementsofthe sculpture and rises forming folds. The perispore thickness ranges from 1to10pm

RESULTS

Thesporesofall thetaxaanalysedaremonolete, duetoitscamerateandfoldedconditions. Then these ellipsoidaltospheroidalinpolar view and planeto valuesvaryaccordingtothepartofthe spore where concave-convexinequatorial view. The shape is only theyareconsidered.

evidentusing the LMsince it is modified by the fea- Noevidence ofamiddlestratumwithpillarswas turesof theouterlayer of thesporoderm (perispore). found in theperispore ofAspleniumpraemorsum.The By thecomparative analysis of the polar and equa- perisporeseemstobecompactatthis level.The typi-torial diameters (Table 1

,

Fig. 1 andFig. 2) it ispossible cal spaces of thisstratumareapparently confinedto todifferentiatethreegroups:1.Sporesupto40pm(A. theinside of the folds in thisspecies.

argentinum,A. depauperatum and A. lorentzii); 2. Inallthe sporesanalysedasupralaesural foldispresent. Spores from 40to50pm (A.achólense,A.formosum, Accordingtotheterminologyproposedby Puttock A.gilliesit] A. laetum,A. lilloanum, A. palmeri, A. &

_Quinn

(1980), thefolds inthe studiedspeciesare: praemorsum,A.pumilum, A.serra,A.triphyllum and 1. winged(withwings):theyare presentin A. argen-A.tucumanense). 3. Spores with diameters higher than tinum(Fig. 3 D-F), A.auritum(Fig. 3 G-I), A. gilliesii 50pm (A. auritum,A. monanthes andA.resiliens). (Fig.4F,H,I),d.laetum(Fig.5A-C),A. lilloanum(Fig. 5D, The laesura is monolete, 16-30pmlong, straight E, G),A. lorentzii(Fig. 5 F.H),A.monanthes (Fig. 5 1, J), and tenuimarginateatthe exospore level.Itisah A.palmeri(fig. 6 A-C),A.pumilum(Fig. 6¥-H),A. resi-waysmarkedontheperispore byacontinuousfold, liens(Fig. 7A-C),A.serra(Fig. 7 D-F) andA.

tucuma-Theexosporeis light-brownand darker than the nense(Fig. 7 J-L); 2. ridged(withridges): theyare perispore in acetolyzedmaterial when observed with presentiñ A. achólense (Fig. 3A-B), A.depauperatum LM.Itis 1.2- 2.5pmthick, single- layered and hasa (Fig.4A-B),A.

_formosum

(Fig.4C-E,G),A.praemorsum

smoothsurface. (Fig. 6 D,E)andA.triphyllum (Fig. 7 G-I).

Table1.Cuantitativedata ofsporesof Asplenium L. from North-West Argentina. Dimensions (in pm)arepresentedinthe form:

minimun (mean) maximum

EQUATORIAL DIAMETER (MINOR) EQUATORIAL DIAMETER (MAJOR) PERISPORE EXOSPORE THICKNES POLAR DIAMETER LAESURA LENGHT SPECIES folds thickness

3.6- 6.0 31.5 (46.5)51.3 31.5 (39.0) 52.2 40.5 (47.9)*51.3 18.0 (30.3) 36.0 1.1 (1.5)1.9 0.4 (0.6) 0.7

A. achalense

23.4 (27.9) 32.6 22.5 (21.5) 26.9 24.8 (27.4)31.9 14.2 (16.3) 22.7 0.4(0.7)1.0 0.3 (0.4) 0.6 3.2- 5.2 A.argentinum

0.5 (0.7) 1.0 0.4 (0.5) 0.6 2.0- 5.3 31.5 (34.2) 36.9 42.3 (51.3) 58.5 26.1 (29.9)32.4

A.auritum 43.2 (50.4) 58.5

4.0- 8.0 17.5 (19.5) 22.5 1.2(1.8)2.1 0.4 (0.6) 0.7

35.4 (38.6)41.6 27.6 (31.7) 35.4 32.5(37.2)41.2 A.depauperatum

1.2 (1.9) 2.5 0.4 (0.5) 0.6 2.5- 5.0 27.5 (36.4)46.2 40.0 (47.3) 52.5 18.7 (23.1)34.5

37.5 (47.1)55.0 A. formosum

3.7- 6.2 21.9(27.8)31.2 1.5 (1.9) 2.5 0.4 (0.6) 0.8

35.0 (40.8)44.4 43.1 (47.1)50.6 43.7 (47.7) 50.6

A. gilliesii

0.4 (0.5) 0.6 5.0- 7.5

41.2 (45.1) 50.0 25.0 (26.4)27.5 1.8 (1.9) 2.5 40.0(47.8)51.2 32.5 (36.5) 38.7

A. laetum

0.9 (1.2) 1.8 0.4 (0.8) 1.3 3.6- 7.2 36.0 (41.6) 45.0 43.2 (46.5) 47.7 18.0(22.4) 27.0

45.0 (48.0)54.0 A.lilloanum

A. lorentzii 30.8 (33.4) 36.6 27.1 (29.2) 33.3 27.9(31.9)35.4 15.0(17.8)22.1 1.2 (1.5) 1.8 0.3 (0.5) 0.7 3.7- 6.2 0.3 (0.4) 0.6 2.5- 7.5 43.1 (50.1)53.1 21.9(30.6)37.5 1.2 (1.5) 1.8

43.7 (47.1)51.9 35.0 (41.0) 46.9 A. monanthes

18.0(19.8) 22.1 0.7 (0.9)1!5 0.3 (0.4)0.6 3.6- 5.4 29.7 (33.5) 36.0 40.5 (42.0)42.3

36.9 (43.9) 50.4 A.palmeri

0.6 (0.7) 1.0 1.0- 1.5 21.6 (26.1)31.5 1.1 (1.0)1.5

•42.5 (45.5) 49.7 31.5(34.8)38.8 43.4 (46.5)51.5 A.praemorsum

18.0 (25.9) 27.5 1.8 (2.2) 2.5 0.4(0.5)0.8 6.2-10.0 37.5 (35.8)47.5 33.7(42.6)51.2

34.0(45.9)55.0 A. pumilum

5.0-10.0

26.3(29.0)31.6 1.8 (2.2) 2.5 0.4(0.6)0.8 52.5 (56.6) 60.3 40.8 (45.9) 52.9 50.4 (56.1)60.8

A. resiliens

0.9 (1.5) 1.8 6.2- 8.7 38.7 (44.4) 49.4 19.4 (21.4) 23.1 1.5(1.8)2.1

41.2 (44.5)48.1 27.5(32.1)36.9 A. serra

42.5 (47.3)5Z5~ 43.3 (48.9) 525

1.8 (2.1) 2.5 0.3 (0.5) 0.8 2.5- 5.0 31.2 (37.7) 43.7 21.2(25.3) 30.0

43.7 (50.4)51.2 A. triphyllum

0.3 (0.6) 0.8 5.0-11.2 21.2 (26.9) 30.4 1.2 (1.4) 1.8

40.8 (47.5) 53.7 33.3(38.9)43.3 A. tucumanense

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G. E.Giudiceet

65 60 55 50 45

o

¡Ü 40

1

<0

3 ₃₅

\

t

30 25 20

//////////////ÿy/

**

v*

Species

Fig.1.Comparativemayorequatorial diameters(pm)inspores of Asplenium speciesfromNorth

_-

WestArgentina

Variationswereobserved withrespecttothecon- be recognized: 1.Reticulate,with large and numer-tourinspores withridgedfolds. Theycanbenarrow ousperforationsresultingthus inan outer stratum likein A. achólense, A. depauperatum and A. totallydiscontinuous(A.serra,Fig. 7 D- F ). 2.

Fe-formosum

orbroadlike in A.praemorsumand A. nestrate,withmiddle-sized perforations,located

between folds (A. auritum,Fig.3 G-H and A.

The foldscanbeisolated,

partially

ortotally mônanthes,Fig.5I,'J) resulting inanouter stratum

fused determininga patternwhich isuniqueineach partiallydiscontinuous. 3.Scarcely perforate (in mostof the species), withafewperforationsof

vari-'The foldmargin isechinulate,irregularor smooth, able size,onthe fold marginorbetween folds and

Mostofthe studiedspecies have echinulated folds whe- associatedwith the elements of the sculpture (A. reasthe foldsinA.depauperatum,A.

formosum,

A.auri- gilliesii, Fig.4I; A.lorentzii, Fig.5F).

turnand A. lilloanumareirregular. The folds witha

smoothmarginarepresentin

Asplenium

achólense(Fig. 3 tae,rugulaeandridges which show variation in den¬ sity and locationevenwithin the same specimen. Accordingto the degree of perforation of the These elementscanbedistributed uniformlyinthe outer stratumoftheperispore (Po), threetypes can whole surfaceorrestrictedtocertainareassuchas triphyllum.

species (Table 2).

Theelementsofthesculptureareechinulae,

cris-A,B)andA.praemorsum(Fig. 6D,E).

65 60

t

55

"

t

40 1

I

t

5 35

I

30 25 20

//////////////<//

Species

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Bol.Soc. Argent.

Table2.Characterstateofperisporeand plant habit ofAspleniumspecies from North-WestArgentina

PERISPORE

PLANT HABIT

SPECIES _FOLDS

PERFORATIONS SCULPTURE Margin Density Fusion

Type

epiphyte, epilithic

few rugulae

few total

A. achúlense Ridged smooth

rugulae, cristae

andechinulae terrestrial few total

A. argentinum winged echinulate many

fenestrate

epiphyte,epilithic

few total rugulae

A. auritum winged irregular

terrestrial,epilithic

partial few rugulae

A. depauperatum ridged irregular few

few rugulae epiphyte,epilithic

few partial

A. formosum ridged irregular

rugulae epilithic partial

A. gilliesii winged echinulate many many

cristae,folds

and echinulae terrestrial few partial few

A.laetum winged echinulate

cristae,rugulae

andechinulae epilithic few

winged irregular partial

A. Hlloanum many

epilithic, terrestrial

rugulae A. lorentzii winged echinulate many partial many

Rugulae and echinulae

fenestrate _epilithic echinulate partial

A. monanthes winged many

few echinulae epilithic

irregular few total A. palmeri winged

epiphyte,epilithic few rugulae

ridged smooth total

A. praemorsum many

Echinulae

and cristae terrestrial echinulate total

A. pumilum winged many many

few epilithic

partial rugulae A. resiliens winged echinulate many

reticulate '_rugulae _epiphyte,_epilithic

echinulate few partial A. serra winged

Echinulae and baculae

few epilithic

partial

A. triphyllum ridged echinulate many

rugulae

and echinulae • terrestrial

few A. tucumanense winged echinulate many total

fold margins orbetween folds. Insome cases the werequotedbyPerezRaya etal. (1986) in species of echinulae fuseattheir basesorare,situated onshort Asplenium of Andalucía.

Palynological studies done in Asplenium like ridges (Fig. 5 B-C;Fig. 6 G-FI). The sculptural elements

areclearlyobserved in A.argentinum(Fig.3D-E ), thoseofViane & Van Cotthem (1977), Puttock & A. gilliesii (Fig. 4 F, H ), A. laetum (Fig. 5A-C),

Quinn

(1980),Pangua&Prada(1988),Pradaetal.

A'

palmeri (Fig. 6 A- C), A. pumilum (Fig. 6 F-H ), (1989) and Johns (2000),consideredgroupsbased A.triphyllum (Fig. 71) and A.serra(Fig. 7 D- F). ondifferentcharacteristics always related with the Theperispore character states and theecology spore surface. In thiswaythe groups dependedon of the analysedtaxaaresummarisedinTable 2. The the characteristicsorgroups of characteristics con-taxaanalysed in this workgrowinadiversity ofhab- sideredas important by each author. Difficulties its, suchasepiphytic, epilithicorterrestrial (Table aroseatthe time ofmaking the comparative analysis 2). Nocorrelationwasobserved between theecol- orwhen generalising.

ogyandperispore complexity. Generalcoincidenceswerefoundbetween the spore

characteristics of the material fromNorth-WestArgen¬ tina and those mentioned in previous studieson some ofthesespecies,exceptforsome aspectsgivenby Prada

etal. (1989) for spores of A.

_formosum

and A. •

Thelargestsize recorded for spores of Asple- praemorsum.Thoseaspects werethepresenceofsmall nium auritum

,

A. monanthes andA. resiliens coúld wrinkles in A.praemorsumand the absence ofperfora-be relatedwithahigher ploidy levelsince according tionsin A.

_formosum.

This differences inouropinion

toLoveetal.(1977) and Tryon&Tryon (1982), the arenotrelevantsince thepresenceofsuch

characteris-two firstspeciesaretetraploids and the last oneis tics donotchange the mainpatternofthesporeand anhexaploid. Spores of large size related withan could be due,atleast inpart,tomodifications and increase in the ploidy level and with theepilithic habit limitations determined by dehydration.

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Fig.3. A-C:A. achalense( MP 3733)A:Spore inequatorial view with fused ridged foldswith asmoothmargin. B: detail of the surface that shows ridged foldswithasmooth marginandrugulateareabetween folds.C: aperispore ffactufe thatexposesits threestrata:innerstratum(Pi), middlestratum (Pm)with

concrescentrodlets(arrow)andouter stratum(Po)apparently composed by twosubstrata(arrowhead). D-F:A. argentinum (MP3743),D: sporein equatorial view withafew fusedwinged folds withanechinulated margin determining polygonalareas,E: detailofthesurfaceshowing baculiformor

spiniform processes and perforationsonand between folds;margin of folds is echinulate.F: aperispore fracture thatshows its threestrata;theouterstratum (Po)is echinulateand twoechinulaearefusedattheir bases(arrow);thecameraeareevidentin the middlestratum(Pm)and the innerstratum(Pi)isthinand continuous.G-I: A.auritum(MP3613)G:aspore inequatorial view with winged folds with scarcely echinulate margins. Theareabetween folds is fenestrate. H: detail ofthesurface showing folds with irregular margins andareabetween folds is fenestrate. I:aperispore fracture that shows its threestrata; theinnerstratum (Pi)isthin and continuous; the middlestratum(Pm)has radial rodlets,someofthemarefusedatdifferent levels and theouter stratum (Po) is perforated between folds. Therearealsosome conesand echinulaeonthe folds margins. E: exospore, P: perispore.Bars: 10 pm.

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Bol. Soc. Argent. Bot. (3-4)

A B C

9

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Fig. 4. A-B:/í.depauperatum(MP3616).A: sporeinequatorial view with ridged foldswith anirregular margin and partially fused. B: detail of the surfaceshowing foldswith anirregular margin andareabetween folds isrugulate. C, D, E, G: A.formosum(MP3770) C: sporeinequatorial view with ridged folds partially fused. D:sporeinproximal view. An evident supralaesural foldis present.E: detail of the surfacewith folds withanirregular margin and with rugulae, few echinulae andcones.G:aperisporefracturethatexposesits threestrata:innerstratum (Pi),middlestratum (Pm) withsinglerodlets

( arrow)andouter stratum (Po)which rises forming folds. F, H,I: A. gilliesii ( MP3764)F: distal viewofaspore with winged folds partially fused. Perforationsarealsopresent.H: detail ofthesurfacethat shows foldswithanechinulatemargin andperforationsonthe whole surface. I:aperispore fracturethat showsaperforateouterstratum(Po) which rises formingfolds,amiddlestratum (Pm)withsingle rodlets (arrow) andaninnerstratum (Pi)

(9)

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F-Fig. 5. A-C: A.laetum (MP3763)A: sporeinequatorial view with winged folds which are fused.Theareabetween folds issculptured. B: detail of the surfacethatshows foldswithanechinulatemargin. The foldsarefused formingareticulum. Echinulae and cristaeare evidentin theareabetween folds.

C:detail of cristae and echinulae inthe areabetweenfolds. D, E,G:A.lilloanum.{MP3783)D:sporeinequatorialview with winged.folds, partially fused. E: detail of the surface showingfoldswith anirregular margin and surface between foldswithrugulae, echinulae and cristae G:aperispore fracture that shows theperispore with its threestrata(*Po,Pm,Pi)and the exospore(E). Pi is detached from theexosporeand ispointed byan arrow.F,H: A. lorentzii

(MP 3627).F: distal view ofasporewith abundant, winged folds with echinulate margin.H:detail of the surface betweenfolds with rugulae and perforations. I,J:A. monanthes(MP3746).I: ecuatorialviewofasporewith abundantpartially fused winged foldsándwithan areabetween folds which is fenestrate..J:detail ofthe surface with fold withanechinulate margin and surface between foldswithrugulaeandechinulae.Bars: 10 pm.

(10)

I

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Fig.6. A-C: A. palmen ( MP 3860). A: Spore in proximalviewwithwingedfolds. A supralesural fold is evident. B: distal view. The foldshaveanirregular marginand are fused determining irregular areas. The surface betweenfoldshas echinulae fusedat

their bases. C: detailofthesurfacebetweenfoldswithechinulaeandrugulae. D-E: A.praemorsum(MP 3622).D: spore in equatorial viewwithridged,fused folds. E: detail of the surfaceshowingfoldswithasmoothmargin and rugulae between folds. In some casesaverrucais evident in thecentreofthe areola. F-H: A.pumilum(MP 3760). F: sporeindistal view withwinged foldspartially fused. G and H: detailofthe surface that shows folds with echinulate margins ( arrow) and area between folds withechinulae,cristae (arrowhead) andperforations.Bars: 10 pm.

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c

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Fig. 7.A-C:A resiliens(MP 3736).A:sporeinequatorial view with wingednarrowfolds. B:detailof the surface showing folds withanirregular margin and surface between folds. C:aperispore fractureat theplacewhere afoldrises that shows its threestrata.The continuousouter stratum ( Po)is risen forming folds; the middlestratum (Pm)iscamerateandthe innerstratum(markedbyan arrowhead))isthin, continuous and attachedto theexospore(E).

D-F:A.serra (MP 3740).D: sporein equatorial view with winged folds and reticulate surface. E: detail ofthesurface.The outerstratum(Po)is discontinuous,completely perforatedandfolded.The foldmargin is echinulated. The middlestratumhasevidentdigitaterodlets(arrow). F:aperispore fracture thatexposesitsstrata: outer stratum (Po) withlargeperforations, middlestratum(Pm) with rodlets, baculae and echinuláe. The bases of which

are onthe innerstratum (Pi).E: exospore. G-I:A.triphyllum(MP 458). G: sporeinproximal view with ridged folds. A supralesural fold is evident. H. distal viewwith manyshort foldspartially fused.I: detail of the surface with folds with echinulate margins and echinulae and baculae in theareabetween folds.J-L:A.tucumanense(MP 3585). J: sporein distal viewwithwinged,narrowand fused folds. K:detailof the surface with foldswith anechinulated margin and rugulaeandechinulaein theareabetween folds. L:aperisporefracturethat exposes its threestrata: outerstratum (Po),middlestratum with rodlets(star) whose basesare onthe innerstratum (Pi).Bars:10 pm.

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Therearealsoafew studies doneonsporoderm

_-

foldswithanirregular margin: presentin ultrastrücturewith TEMandSEMonthis genus (Pettitt, A. depauperatum and A.

_formosum

1966;Lugardon, 1972,1974;Tiyon&Lugardon, 1991).

-

foldswithanechinulatemargin: present in Mostofthe Aspleniumspeciesstudiedand ob- A. triphyllum

servedinsection withLM and SEMhaveacomplex Thusseven outofseventeenspeciesstudied could perisporewhich ischaracterisedby having spaces be determinedbytheirspores, theyareA.achalense, with radialrodlets(pillars)inits middlepart.In A. A.auritum,A. monanthes,A.praemorsum, A. resiliens, praemorsum the spacesare locatedand confined A.serraand A.triphyllum.

tothe inside of the folds andnorodlets (or pillars) The groups obtainedbasedonspore character-wereobserved neither in the middlestratumnorin istics,donotmatch withthesystematic groupsbased the inside of folds.It means that spaces in the onvegetative characteristics established by De la spores of thisspeciesarerestrictedtoareas;how- Sota(1977),Tryon&Tryon(1982)andMurakami&

everintherestofthestudied speciesacontinuous Moran(1993).Nevertheless,itshouldbe taken into space exists between thetwostrata( Piand Po)of accountthat theseclassificationsweremade with¬

outusingpalynologicalcharacteristics.

Wewouldliketohighlight that in thecaseof A.

spores of Asplenium trichomanes in different serrathereisacongruencebetweenexo

-morpho-stagesofdevelopment indicate that the featuresof logicalcharacteristics (bladearchitecture,sizeand perisporeouter stratum areformedduring latesteps, pinnaemargins) and spore characteristics

which

We consider that studies onsporoderm develop- makes thesespecieseasilydetermined in theareaof mentand hltrastructure wouldprovideusefiildata study.

about morphologyrelatedtofunctionand thatthey theperispore.

Observationsmade byTigerschiõld(1981)on

Aswehavepreviouslypointedoutnot correla-would allowustosearch forsimilaritieswith other tionwasfoundbetweentheperispore surfacecom¬ plexity and the ecologyinthe studiedspecies,since Filicopsids.

Asaresultof thisanalysisthe studiedspecies weobservedinepiphyticspeciesslightlyornamented canbe classifiedintotwogroupsaccordingtothe

-

perforatespecies (A. achalense)torugulate- slightly perforate (A.

formosum,

A. praemorsum) tofenes¬

trate(A.auritum)totheextremecaseofthosewith completelyperforate (reticulate)surface inA.serra. A direct correlationbetweenthecomplexity ofthe

-Perisporefenestrate: A. monanthes and perispore surfaceand the ecologywas quoted by

Tryon(1990)basedonthe analysis of 111speciesof charactersstatesoftheperispore surface:

1-Perispore with winged folds -Perispore reticulate: A.serra

A.auritum

-Perisporewithafew perforations: with varied Asplenium. ornamentation,like:echinulae,cristae and rugulae

located in theareabetween folds: A. argentinum, paragraphs,weconsider that spore characteristics A.gilliesii,A. lilloanum, A. lorentzii, A.palmeri, should be taken into account together with other A. resiliens andA.tucumanensè.Itismoreevident morphological featuresandthattheycould help in inA.pumilum and A. laetum. It is difficulttodiffer- thedeterminationofmost taxaofAsplenium of North-entiate thesespeciesby their spores duetovaria- WestArgentina. Inthenextcontribution(Morbelli tionsin ornamentation,densityofthesculpturalele- etal.,in prep.), thesporodermultrastructurewill be

mentsandcharacteristicsofthe folds margins. Nev- . analysedunder scanning and transmission electron ertheless,A. resiliens could be distinguished from

.

microscopyinthe structuralpatternsrecognised in therestbytheir large spores. In the sameway,A. this study.

laetum and A.pumilumcanbe distinguished from therestby their foldswith markedlyechinulated

marginsand

the.

surfacebetween folds with cristae

ACKNOWLEDGMENTS

together withfusedechinulae formingareticulum.

Accordingto the concepts- given in previous

The authors thank RafaelUrrejola

of

"Servicio de Microscopía Electrónica del Museo de Ciencias 2.Perisporewithridged folds

-

foldswitha smoothmargin: present' in _Naturales_{de La Plata". This work}_was_supported bya grantfromthe National Councilof Scientific A. achalense andA. praemorsum

(13)

G.E.Giudiceet al.,Palynological study in

NAYAR, B. K., LATA, P & TIWARI, P. 1964. Sporesmor¬

phology oftheferns ofwest TropicalAfrica. Pollenet

Spores6: 545-582.

PANGUA, E. & C. PRADA. 1988.Tipos esporales en

Aspleniaceas Ibéricas.Lagascalia 15:157-167. PEREZRAYA, F., CASARES PORCEL, M. , MOLERO

MESA, J. & GONZALEZ TEJERO, M. R.. 1986. Estudio

palinológicodel géneroAspleniumL. En Sierra Nevada (Andalucía- España). Ccmdollea41:368-380. PETTITT, J.M. 1966.Exine structureinsome fossiland

recent sporesand pollenasrevealed by light and elec¬

tronmicroscopy. Bull. Brit.Mus.. Geol.13: 223-257. PONCE, M. 1996. Pteridophyta. In ZULOAGA, F. & O.

MORRONE (eds.), Cátalogo de las Plantas Vasculares de la Rep.ArgentinaI.Monogr.Syst. Bot. Missouri Bol. Gard. 60: 1-79.

PRADA,C., PANGUA,E., BLANCO, P., CUBAS, P. & C. PARDO. 1989. LasAspleniaceasde los Herbarios de MutiseIsern. Anales Jará. Bot. Madrid 46: 539-552. PUTTOCK, C. F. &QUINN,C.J. 1980.Perisporemor¬

phologyand thetaxonomyof AustralianAspleniaceae.

Austr. J. Bot. 28: 305-322.

SOTA, DELA, E.R. 1977. Pteridophyta. Ih: A. CABRERA (ed.) Flora de la Provincia de Jujuy. Colecc. Ci. INTA 13: 1-275 TIGERSCHIÕLD, E. 1981. TheAspleniumtrichomanes

complex inEastCentral Sweden. Nord. J. Bot. 1: 12-16. TRYON, A. F.1990. Fern spores: evolutionary levels and

ecologicaldifferentiation.PI.Syst.Evol. 5:71-79. TRYONA.F & LUGARDON, B. 1991.Spores _ofthe

Pteridophyta. Springer-Verlag, New York.

TRYON,R.M. & TRYON, A.F. 1982. Ferns and Allied

plants.Spinger-Verlag,New York.

VIANE, R.H.& VAN COTTHEM,W. 1977. Sporemorphol¬

ogy and stomatal characters ofsomeKenyanAsplenium species.Ber. Deutsch.Bot.Ges.90:219-339.

andTechnologicalResearch (CONICET),Buenos

Aires,for PIP 5044 anda grantfromthe National University ofLaPlata forProject N°363.

BIBLIOGRAPHY

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(Pteridophyta) de laprovinciade Buenos Aires, Argen¬ tina.Darwiniana 32:131-137.

MORTON,C.V. & D. LELLINGER. 1966. The

Polypo-diaceae subfamilyAsplenioideaein Venezuela. Mem. New YorkBot.Gard. 15:1-49.

MORBELLI, M.A. 1980. Morfología de las esporas de las Pteridophytapresentes enlaRegiónFuego-Patagónica. RepúblicaArgentina.Opera Lilloana 28: 1-138.

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(Aspleniaceae). Ann. Missouri Bot. Gard. 80: 1-38. NAYAR,B. K. & S. DEVI. 1964. Sporemorphology of

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Recibido el 12 de Abril de 2002, aceptado el 04 Septiembre de2002.