Morphological diversity of ferns in the Dryopteris affinis group in ...

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Morphological diversity of ferns in the Dryopteris affinis group in Central Poland BEATA WOZIWODA Department of Geobotany and Plant Ecology, University of £ód, Banacha 12/16, 90-237 £ód; e-mail: [email protected] ABSTRACT : The article presents the morphological diversity of individuals from Dryopteris affinis group, which grow in 8 isolated localities in the £aska Upland (Central Poland). It contains descriptions and comparison of 17 morphological features that are often used as diagnostic in manuals. The studied ferns represent two species: D. borreri (Newm.) Oberh. & Tavel and Dryopteris cambriensis (Fraser-Jenk.) Beitel & W.R. Buck, which means that D. affinis (Löwe) Fraser-Jenk. s. s. has not been found in this area so far. ABSTRAKT : Praca przedstawia zmiennoæ morfologiczn¹ okazów z grupy Dryopteris affinis, rosn¹cych na 8 izolowanych stanowiskach na Wysoczynie £askiej (Centralna Polska). Zawiera opis i porównanie 17 cech morfologicznych wykorzystywanych w kluczach. Badane paprocie nale¿¹ do dwóch gatunków: D. borreri (Newm.) Oberh. & Tavel i Dryopteris cambriensis (Fraser-Jenk.) Beitel & W.R. Buck, co oznacza, ¿e D. affinis (Löwe) Fraser-Jenk. s. s. nie zosta³a dotychczas na tym terenie odnaleziona. KEY WORDS: ferns, Dryopteris affinis agg., Dryopteris borreri, Dryopteris cambriensis, morphology, Central Poland.

Introduction The Scaly Male-fern Dryopteris affinis group is one of the most interesting and difficult objects of taxonomic studies in the European pteridophyte flora (Pigott 1997; Ekrt et al. 2009). This taxon reproduces without the necessity of fertilization (Pigott 1997). The plants are apomictic that means they can generate vegetative clones, and each local population can be regarded as a new species (Whild, Lockton 1999). Moreover, they can easily form pure strains

WOZIWODA B. 2009. Morphological diversity of ferns in the Dryopteris affinis group in Central Poland. In: E. Szczêniak, E. Gola (eds), Genus Dryopteris Adans. in Poland. Polish Botanical Society & Institute of Plant Biology, University of Wroc³aw, Wroc³aw, p. 4559.

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and partially fertile hybrids with D. filix-mas (L.) Schott and ferns from D. carthusiana group (Piêko-Mirkowa 1981; Pigott 1997). In addition, the variable environmental conditions can result in significant and often confusing changes of fern morphology (Piggot 1997). There have been published numerous approaches to the D. affinis complex treatment over the years, with changing classifications of species, subspecies, varieties and morphotypes (Pigott 1997; Whild, Lockton 1999). The newest description of the D. affinis group is presented by Fraser-Jenkins (2007). In Central Europe, three taxa differing in the ploidy level and evolutionary history are currently recognized (Fraser-Jenkins 2007; Ekrt et al. 2009). Diploid 2n=82 D. affinis (Löwe) Fraser-Jenk. s.s. is restricted to the western and southern parts of Central Europe. Two triploid 2n=123 species, D. borreri (Newm.) Oberh. & Tavel and D. cambrensis (Fraser-Jenk.) Beitel & W.R. Buck, are known from Central Europe. Furthermore, in continental Europe three other triploid taxa are noted: D. pseudo-disjuncta (Tavel ex FraserJenk.) Fraser-Jenk. in Western Europe, D. schorapenensis Askerov and D. pontica Fraser-Jenk. in the surroundings of the Caucasus Mts (FraserJenkins 2007). The variety of Dryopteris affinis group in Poland is not recognized yet; only a few information about occurrence of D. cambrensis and D. borreri are known (Fraser-Jenkins 2007; Ekrt et al. 2009). The aim of presented research was to determine the taxonomical position of specimens occurring on scattered and isolated stands in the £aska Upland as well as the range of their morphological diversity.

Characteristics of Dryopteris affinis aggregation Distribution: Dryopteris affinis agg. occurs in Europe, northern Africa, Macaronesia and south-western Asia (the Caucasus Mts). The distribution of this taxon in the western part of Europe is related to the Atlantic climate. The fern is the most abundant in areas with high humidity such as the British Isles and western France (Watson, Dallwitz 2004; Fraser-Jenkins 2007; Byrne et al. 2008). In the continental part of Europe, it is considered as a mountain species (Fraser-Jenkins 2007). In Poland D. affinis is distributed mainly in the Carpathian Mts (PiêkoMirkowa 1981), where it occurs in the sub-mountain and lower mountain zones up to an altitude of 1100 m above sea level. It grows in coniferous forests (with Abies alba and Picea abies) and in beech woods; also in shady places among rocks and streams, mostly on acidic soils (Piêko-Mirkowa, Miechówka 1992; Piêko-Mirkowa et al. 1999). Scaly-male fern has also been reported from

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dispersed localities in Polish lowlands: in Góry wiêtokrzyskie Mts, in the Upper Silesia Province, in the Wy¿yna Lubelska upland, near Toruñ, £ód (£aska Upland), as well as in the Mazurian Lake District (Piêko-Mirkowa 1981; Mirek 1995; Zaj¹c and Zaj¹c 2001; Woziwoda 2006; Podsiedlik 2009; Tla³ka 2009). Morphology: according to Piêko-Mirkowa (1981), Fraser-Jenkins (1993, 2007) and Watson and Dallwitz (2004). Leaves of Dryopteris affinis are tufted at the crown, (30) 50-90 (160) cm long. In early summer, the fronds are yellowish-green and become darker later in the year. They are persistent or dying out progressively through winter. Fronds are the widest in the middle and narrowing towards the apex, with more or less glossy and stiff lamina. The petiole is shorter than the blade, usually the .1 5 -1 6 up to the ¼ of the blade length. The stipe is particularly strong and stout, covered with golden-brown or light orange-brown glossy narrowly lanceolate scales. Dense hair-like scales (fibrillae) are also present on the rachis and are often found on the lamina surface. Leaves are compound and divided, with visibly divided pinnae. The pinnae length is decreasing gradually towards the base of the blade; the lowest ones are relatively short, the longest pinnae are in the middle of the blade. The lowermost pinnae pair is up to half the length of those in the central part of the frond. At the junction point of each pinnae with the rachis, the dark pigment forms a distinctive black patch or spot around the pinna-midrib base (! sometimes visible only on the fresh leaves; patches can disappear during drying of the plant material). The shape of pinna-segments is very characteristic: it makes rectangular lobes, with square- or rounded-truncated apex, sometimes with acute notches only at the apex. This fern has no clear distinction between fertile and sterile (vegetative) leaves. Sporangia are superficial and gathered in sori; sori are more or less circular in outline, usually smaller than in D. filix-mas. They remain covered by indusium. The indusium is stiff, with edges covering sporangia. It is persistent and usually remains on overwintered fronds.

Materials and methods Plants used for the study came from all localities of D. affiinis agg. known in the £aska Upland. Examined 8 stands of ferns are isolated and very small only 1 or 2 individuals are noted in each of them (Woziwoda 2006). Used symbols of stands and detailed localizations are presented in Table 1. The 17 morphological characteristic features of individuals are described: lamina shape, lamina base, lamina texture, frond persistence, pinna shape in the mid blade, pinna-segment (= pinnulae) shape, pinna-segment apex, the lowest

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pair of pinna-segment on each pinna, the lowest basiscopic pinna-segment on the lowest pinnae of frond, pinna/pinna spacing; pinna-segment/pinna-segment spacing, pinna-midrib base, stipe thickness, scale colour, scale shape, scale density and indusium shape. Morphological description of features in analysed specimens, named with letters from A to H, is summarised in Table 1. The sample is too scarce to make statistical analyses. The classification of ferns and nomenclature of D. affinis group follow FraserJenkins (2007). A list of localities is given in ATPOL grid squares 1 × 1 km (Zaj¹c 1978) and forest divisions. Herbarium sheets of plants are deposited in the LOD (Herbarium Universitatis Lodziensis).

Results The study of morphological characteristics of plants from the £aska Upland lead us to two species: Dryopteris cambriensis (Fraser-Jenk.) Beitel & W.R. Buck (Tab.1: A; Figs 1, 3, 5: A) the specimen from Dobków-Julianów, and Dryopteris borreri (Newm.) Oberh. & Tavel (Tab.1: B-H; Figs 1-5: B-H) occurring in next seven stands. Only three of the analysed features distinguish D. cambrensis: narrow lamina, distinctly acute dentate and glandular pinnulae apex, and the fact that fronds die out after first frost. Respectively, laminas of D. borreri are wider, pinnulae apex has slightly acute to obtuse teeth or it is without teeth, and fronds die out progressively through winter in mild winter ferns are semi-evergreen. The range of variation of other analysed features is similar in both species. All ferns have fronds which arise irregularly from clumps. The young fronds are light green, lighter than leaves of other Dryopteris species, which grow in the neighbourhood, then they change the colour to dark green. Leaf blades are lanceolate to oblong or to ovate with tapering-truncate or rarely truncate base (Figs 1-2), slightly glossy, stiff, slightly stiff or flexible (Tab. 1: Lt). The lamina length is about 35-45 cm (33 to 60, average 40 cm). Pinnae: the pinna-midrib base has usually a distinctive black spot or black patch, which mostly disappears during drying of herbarium specimens. In specimens G and H, black patches are slightly distinguished (Tab. 1: Pm). Pinnae are oblong, oblong-triangular to triangular (Figs 2-3), having in the midblade about 10 cm in length; their tips are tapered to acuminate. Pinnulae are usually squarely-truncate at the base of pinnae, rounded-truncate in the middle part, to slightly oblique-truncate at the top, without or with slightly or obtuse acute teeth. In the truncate-pinnuled parts of pinnae, the teeth are usually longer than in the apex center and protrude above the corner of each pinna-segment. The lower pinnulae on lower pinnae, particularly the lowest basiscopic one of

P-sa

Lsh

D oblong to ovate (Fig. 1D); 34 cm long tapering-truncate

E F lanceolate to reverse lanceolate to oblong ovate (Fig. 2E); (Fig. 2F); 36 cm long (36)-54 cm long tapering-truncate truncate or slightly tapering-truncate glossy, stiff slightly glossy, glossy, slightly not stiff stiff dying out dying out dying out progressively progressively progressively through winter through winter through winter with a distinctive with a distinctive with a distinctive black spot black spot black patch oblong triangular, narrowly triangular, oblong, tapering to tapering to the acute tapering to shortly acuminate tip tip (?- most of pinna acuminate tip (Fig. 4F1, F2); tips are damaged by (Fig. 4E1, E2); (9,5-)11,5-12 cm insects or frost) 7,5-9,5 cm long long (Fig. 3D1) even, with parallel even, strongly even, rectangular rectangular lobed, lobed, most of them sides, rectangular lobed (Fig. 5F) with slightly pointed slightly tapering to side-lobes and their acute tip parallel sides (Fig. 5E) (Fig. 3D1, D2) rounded-truncate squarely-truncate to rounded-truncate to rounded-truncate squarely-truncate squarely-truncate (at or slightly rounded-truncate slightly squarely(looking as pinnae- the base of pinna) squarely-truncate with obtuse teeth or truncate with obtuse segments have been (Fig. 5E), roundedwith numerous without teeth or slightly acute teeth cut with a pair of truncate to slightly distinct acute (Fig. 5B) or without teeth scissors) to slightly oblique-truncate (at teeth (Fig. 5A), oblique-truncate, the top of pinna), glandular with acute scattered with scattered acute teeth, sometimes teeth longer at the corners

A B C narrow, lanceolate lanceolate to oblong lanceolate to oblong to ovate (Fig. 1A); (Fig. 1B); (Fig. 1C); 33 cm long 45 cm long 45 cm long Lb tapering-truncate slightly taperingtruncate truncate Lt slightly glossy, not slightly glossy, slightly glossy, stiff stiff not stiff Fp dying out dying out dying out after first frost progressively progressively through winter through winter Pm with a distinctive with a distinctive with a distinctive black patch black spot black spot Psh triangular, tapering oblong, triangular, oblong, triangular, from middle of tapering to shortly tapering to shortly blade to shortly acuminate with acuminate tip acuminate tip slightly turned up tip (Fig. 3C1, C2); (Fig. 3A1, A2); (Fig. 3B1, B2); 10 cm long 6,0-6,5 cm long 10 cm long P-ssh rather rectangular even, with parallel even, rectangular lobed sides, rectangular lobed (Fig. 5C) lobed (Fig. 5B)

Tab. 1 H lanceolate to ovate (Fig. 2H); 60 cm long tapering-truncate

Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group even, rectangular lobed (Fig. 5H)

rounded-truncate, slightly oblique, with two (or more) acute teeth (Fig. 5H)

even, strongly rectangular lobed, with parallel sides (Fig. 5G)

squarely-truncate (Fig. 5G)

slightly glossy, not stiff dying out dying out progressively progressively through winter through winter with a slightly visible with slightly visible black patch black patch oblong triangular, oblong, triangular, tapering to shortly tapering to acuminate tip acuminate tip (Fig. 4H1, H2); 10 cm long

slightly glossy, stiff

G lanceolate to oblong (Fig. 2G); 38 cm long truncate

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as long as adjacent pinna-segments, stalked only on the acroscopic pinnasegment on the lowest pinnae

robust, 14 cm long reddish gold to dark brown

robust; 22 cm long reddish gold

mostly narrow, both types wide mostly narrow, wide and narrow at the twisted, wide ones at and narrow at the the stipe base stipe base, narrow stipe ones the rachis

Ssh

Sc

Sth

narrow, twisted

light brown to pale yellowish

slender, 12-13 cm

distant (Fig. 5E)

slightly longer than adjacent pinnasegments, stalked only on the lowest pinnae,

robust, 8 cm long

distant in young fronds, than overlapping distant to overlapping

moderately robust, 17 cm long pale straw with a darker base

distant (Fig. 5H)

distant

slightly larger than slightly larger and corresponding ones more developed than on pinnae corresponding ones immediately above, on pinnae partly attached immediately above, partly attached at the base

as long as adjacent pinna-segments, stalked only on the lowest pinnae

reddish gold to light reddish gold with a brown with a darker darker base base mostly narrow, also mostly narrow wide to narrow at the wide ones at the stipe stipe and narrow at base the rachis

slender, (17,5) 27 cm

distant (Fig. 5F)

slightly longer than slightly longer and adjacent pinnamore developed segments, stalked (with a basal auricle) only on the lowest than adjacent pinnapinna segment, asymmetrically attached at the base larger and more slightly more developed than developed than corresponding ones corresponding ones on pinnae on pinnae immediately above, immediately above, lobed with a small lobed with small basal auricle, partly square basal auricles, attached at the base attached at the base to the pinna-midrib distant (Fig. 2E) distant (Fig. 2F)

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distant to slightly separate

slightly separate (Fig. 5C)

slightly separate to slightly separate to overlapping overlapping (Fig. (Fig. 5A) 3A1, B2), pinnulae gathered (Fig. 5B) moderately robust, moderately robust, 13 cm long 22 cm long light brown to pale dark brown yellowish

overlapping (Fig. 3D1)

slightly longer, wider and more deflexed than adjacent pinnasegments, stalked or partly attached at the base from midlamina downwards larger than corresponding ones on pinnae immediately above, lobed with a rectangular side-lobe and a basal auricle, fully stalked

P-s/ P-s

slightly larger and more developed than corresponding ones on pinnae immediately above, partly attached at the base

slightly longer than adjacent pinnasegments, stalked only on acroscopic segments on the lowest pinnae

slightly separate

as long as adjacent pinna-segments, but wider and lobed with a rectangular basal auricle, stalked on pinnae from midlamina downwards slightly larger and more developed than corresponding ones on pinnae immediately above, fully stalked

slightly separate to overlapping

P/P

distant (Fig. 3A1, A2)

Lbp-s slightly larger and more developed than corresponding ones on pinnae immediately above, attached at the base

Lp-s

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moderately dense dense at the stick to scarce at the rachis shriveling and slightly lifted up to a lifted to a distorted clear cone, not cone with wavy splitting inflected indusium margins, not splitting indusia large and thick, only slightly lifted with slightly inflected margins, not splitting

scarce moderately dense

moderately dense

moderately dense

moderately dense

thick, shriveling and shriveling and lifted thick, shriveling and thick, slightly lifted, shriveling and lifted lifted to a cone, not to a distorted cone lifted to a cone, not not splitting to a distorted cone splitting with wavy inflected splitting with wavy inflected indusium margins, indusium margins, not splitting not splitting (Fig. 5E)

very dense (Fig. 3D2)


Gatunki: A – Dryopteris cambriensis; B-H – Dryopteris borreri Lista stanowisk: A – Dobków-Julianów, 225a oddzia³ leny; DD8314 kwadrat ATPOL; B – Dobków-Julianów, 224f-1; DD8314; C – Dobków-Julianów, 224f-2; DD8314; D – Dobków-Julianów, 225b; DD8314; E – Pelagia I, 238l-1; DD8323; F – Pelagia I, 238l-2; DD8323; G – Pruszków, las prywatny; DE0303; H – Rzepiszew 159b; DD8302. Skróty analizowanych cech morfologicznych: Lsh – pokrój blaszki; Lb – podstawa blaszki; Lt – struktura blaszki; Fp – trwa³oæ lici; P-m – nasada odcinków I rzêdu; Psh – pokrój odcinków I rzêdu w rodku blaszki (pinnae); P-ssh – pokrój odcinków II rzêdu (pinnulae); P-sa – wierzcho³ek odcinków II rzêdu; Lp-s – najni¿sza para odcinków II (pinnulae) rzêdu ka¿dego odcinka I rzêdu (pinnae); Lbp-s – najni¿szy odcinek II rzêdu (pinnulae) u podstawy najni¿szego segmentu (pinna); P/P – rozstawienie odcinków I rzêdu; P-s/P-s – rozstawienie odcinków II rzêdu; Sth – gruboæ ogonka; Sc – kolor ³usek; Ssh – pokrój ³usek; Sd – gêstoæ ³usek; Ish – kszta³t zawijki.

The species: A – Dryopteris cambriensis; B-H – Dryopteris borreri List of localities: A – Dobków-Julianów, 225a forest division; DD8314 ATPOL grid square; B – Dobków-Julianów, 224f-1; DD8314; C – Dobków-Julianów, 224f-2; DD8314; D – Dobków-Julianów, 225b; DD8314; E – Pelagia I, 238l-1; DD8323; F – Pelagia I, 238l-2; DD8323; G – Pruszków, private forest; DE0303; H – Rzepiszew 159b; DD8302. Abbreviations of analysed morphological features: Lsh – lamina shape; Lb – lamina base; Lt – lamina texture; Fp – frond persistence; P-m – pinna-midrib base; Psh – pinna shape in the mid blade; P-ssh – pinna-segment (= pinnulae) shape; P-sa – pinna-segment apex; Lp-s – the lowest pair of pinna-segment (pinnulae) on each pinna; Lbp-s – the lowest basiscopic pinna-segment (pinnulae) on the lowest pinna of frond; P/P – pinna/pinna spacing; P-s/P-s – pinna-segment/pinna-segment spacing; Sth – stipe thickness; Sc – scale colour; Ssh – scale shape; Sd – scale density; Ish – indusium shape.

Ish

Sd

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Fig. 1. Fronds. Lamina shape and lamina base. A-D according to the text Ryc. 1. Licie. Kszta³t licia i podstawy blaszki liciowej. A-D zgodnie z tekstem

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Fig. 2. Fronds. Lamina shape and lamina base. E-H according to the text. Ryc. 2. Licie. Kszta³t licia i podstawy blaszki liciowej. E-H zgodnie z tekstem

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Fig. 3. Pinna shapes. A-D according to the text; 1 upper side, 2 lower side. Ryc. 3. Pokrój odcinków I rzêdu (pinnae). A-D zgodnie z tekstem; 1 strona grzbietowa, 2 strona brzuszna

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Fig. 4. Pinna shapes. E-H according to the text; 1 upper side, 2 lower side Ryc. 4. Pokrój odcinków I rzêdu (pinnae). E-H zgodnie z tekstem; 1 strona grzbietowa, 2 strona brzuszna

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Fig. 5. Pinna-segments (pinnulae), scales on rachis and indusia. A-H according to the text Ryc. 5. Pokrój odcinków II rzêdu (pinnulae), ³uski na osi i zawijki. A-H zgodnie z tekstem

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the frond, are usually lobed with a characteristic rectangular side lobe and a rectangular basal auricle, slightly larger and more developed than corresponding ones on pinnae immediately above. The lowest pair of pinnulae on each pinna is as long as adjacent pinnula or slightly longer, with the lowest basiscopic pinnula on the lowest pinna slightly larger or larger than corresponding ones on pinna immediately above. Pinna/pinna and pinnula/pinnula spacing is different: elements are distant, slightly separate to overlapping (Figs 4-6). Stipes: relatively long, moderately robust or robust, only in specimens E and F slender, twice or three times shorter than a lamina. Stipes and rachis are usually clothed with moderately dense, dense or very dense scales; only in the specimen C scales are scarce. Scales: wide and narrow (then wide scales occur at the base of the stipe) or only narrow, in specimens B and E twisted; light brown to reddish gold or dark brown, in specimens F, G and H with darker base (Tab.1: Sc). The pale narrow scales occur on the abaxial surface of a lamina. Indusium: when mature thick, shriveling and lifted to a distorted cone, not splitting, in specimens A, C, E and H with inflected or slightly inflected margins (Tab. 1: Ish; Fig. 5). Between analysed D. borreri specimens, the most distinct are specimens H (not stiff lamina texture, pinnulae apex with a few acute teeth, a black patch at the base of pinnae only slightly visible) and C (lamina texture not stiff and low density of scales).

Discussion and conclusion The results of presented study indicate the presence of Dryopteris borreri and D. cambriensis, and the lack of individuals of Dryopteris affinis s.s. in the studied area. The studied plants of D. borreri and D. cambriensis from localities in the £aska Upland are all known individuals from this area so far. All studied stands are located in the secondary habitats of Pinus sylvestris young unstable communities (Woziwoda 2006). At present, any locality in a natural habitat is not known. Occurrence in habitats that are atypical of these ferns could be one of the reasons of the fern morphological variability observed in studied area. The most interesting are variations of the features accepted as typical of D. affinis aggregation: presence of the distinctive black patch at the base of pinnae, a squarely-truncate top of pinnulae and a stiff texture of leaf blade (Tab. 1, features Pm, P-sa, Lt). Observed modifications suggest that further examination of ferns from Dryopteris affinis group is needed. The fern individuals are isolated, but they grow in clusters with D. filix-mas, D. carthusiana and D. dilatata. Probably each of those species may be a

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male parent in crosses with Scaly male-fern, and potentially with other Dryopteris species. Moreover, the frond morphology of D. ×critica, the hybrid between D. borreri and D. filix-mas, is generally very similar to that of D. borreri (Ekrt et al. 2009), so it is very easy to make a mistake in taxonomic classification. In Britain, Czech Republic and Slovakia, there are examples of individual plants with morphology difficult to classify to a known form. Morphological variability within the D. affinis group is extremely high (FraserJenkins 2007, Byrne et al. 2008). At present four hybrids between the D. affinis agg. are treated at the nothospecific rank and two of the hybrids are divided into six nothosubspecies (Fraser-Jenkins 2007). The ferns from Poland (PiêkoMirkowa 1981; Mirek 1995; Piêko-Mirkowa et al. 1999; Woziwoda 2006; Podsiedlik 2009; Tla³ka 2009) may represent varying degrees of the genetic and genomic differences. The chromosome number determination and the DNA ploidy level estimation are necessary. We ought to study morphological diversity within the Dryopteris affinis group, the range of species, subspecies and varieties, their geographical and ecological distribution in Poland Acknowledgements: The author greatly thanks to Dr. Libor Ekrt for his valuable comments, taxonomic revision of D. affinis agg. herbarium specimens and determination of D. borreri and D. cambriensis.

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Zró¿nicowanie morfologiczne paproci z grupy Dryopteris affinis agg. w Polsce rodkowej Dryopteris affinis agg. wykazuje silne zró¿nicowanie genetyczne i morfologiczne. Aktualnie w obrêbie tego taksonu wyró¿nia siê 6 gatunków z podgatunkami i wariantami oraz mieszañce miêdzygatunkowe. W Polsce gatunek wystêpuje przede wszystkim w Karpatach. Na obszarze Wysoczyzny £askiej (Polska rodkowa) jest bardzo rzadki, znany z zaledwie 8 izolowanych wyst¹pieñ, licz¹cych 1-2 roliny. Okazy tej paproci charakteryzuj¹ siê du¿¹ zmiennoci¹ kszta³tu blaszek liciowych (Ryc. 1, 2), odcinków I i II rzêdu (Ryc. 3, 4, 5), a tak¿e zró¿nicowaniem barwy, koloru i kszta³tu ³usek okrywaj¹cych ogonek i o licia. Istotna jest zmiennoæ cech wskazywanych jako diagnostyczne dla D. affinis agg.: obecnoci wyranej czarnej plamki u nasady listków, sztywnoci blaszki liciowej i gêstoci ³usek na ogonku (Tab. 1 cechy Pm, P-sa, Lt). Zgromadzone dane wskazuj¹ na przynale¿noæ wiêkszoci odnotowanych osobników do gatunku Dryopteris borreri (Newm.) Oberh. & Tavel. Analiza cech morfologicznych okazu z Dobkowa-Julianowa z oddz. 225a potwierdza jego przynale¿noæ do Dryopteris cambriensis (Fraser-Jenk.) Beitel & W.R. Buck. Oba taksony by³y dotychczas zaliczane do D. affinis agg.