Morphological diversity of ferns in the Dryopteris affinis group in

Morphological diversity of ferns in the Dryopteris affinis group in

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Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
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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 £ódŸ, Banacha 12/16,
90-237 £ódŸ; 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 zmiennoœæ morfologiczn¹ okazów z grupy Dryopteris
affinis, rosn¹cych na 8 izolowanych stanowiskach na WysoczyŸnie £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. Szczêœniak, E. Gola (eds), Genus Dryopteris Adans. in Poland. – Polish
Botanical Society & Institute of Plant Biology, University of Wroc³aw, Wroc³aw, p. 45–59.
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46
and partially fertile hybrids with D. filix-mas (L.) Schott and ferns from D.
carthusiana group (Piêkoœ-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 (PiêkoœMirkowa 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 (Piêkoœ-Mirkowa, Miechówka 1992;
Piêkoœ-Mirkowa et al. 1999). Scaly-male fern has also been reported from
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Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
47
dispersed localities in Polish lowlands: in Góry Œwiêtokrzyskie Mts, in the Upper
Silesia Province, in the Wy¿yna Lubelska upland, near Toruñ, £ódŸ (£aska
Upland), as well as in the Mazurian Lake District (Piêkoœ-Mirkowa 1981; Mirek
1995; Zaj¹c and Zaj¹c 2001; Woziwoda 2006; Podsiedlik 2009; Tla³ka 2009).
Morphology: according to Piêkoœ-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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narrow
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
50
50
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)
Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
Gatunki: A – Dryopteris cambriensis; B-H – Dryopteris borreri
Lista stanowisk: A – Dobków-Julianów, 225a oddzia³ leœny; 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 – trwa³oœæ liœci; P-m – nasada odcinków I rzêdu;
Psh – pokrój odcinków I rzêdu w œrodku 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 – gruboœæ ogonka; Sc – kolor ³usek; Ssh – pokrój ³usek; Sd – gêstoœæ 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. Liœcie. Kszta³t liœcia i podstawy blaszki liœciowej. A-D – zgodnie z tekstem
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Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
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Fig. 2. Fronds. Lamina shape and lamina base. E-H – according to the text.
Ryc. 2. Liœcie. Kszta³t liœcia i podstawy blaszki liœciowej. 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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Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
55
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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Morphological-diversity-of-ferns-in-the-Dryopteris-affinis-group
57
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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58
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 (PiêkoœMirkowa 1981; Mirek 1995; Piêkoœ-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 œrodkowej
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 œrodkowa) jest bardzo rzadki, znany z zaledwie 8 izolowanych
wyst¹pieñ, licz¹cych 1-2 roœliny. Okazy tej paproci charakteryzuj¹ siê du¿¹ zmiennoœci¹
kszta³tu blaszek liœciowych (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 oœ liœcia.
Istotna jest zmiennoϾ cech wskazywanych jako diagnostyczne dla D. affinis agg.:
obecnoœci wyraŸnej czarnej plamki u nasady listków, sztywnoœci blaszki liœciowej
i gêstoœci ³usek na ogonku (Tab. 1 cechy Pm, P-sa, Lt).
Zgromadzone dane wskazuj¹ na przynale¿noœæ wiêkszoœci odnotowanych
osobników do gatunku Dryopteris borreri (Newm.) Oberh. & Tavel. Analiza cech
morfologicznych okazu z Dobkowa-Julianowa z oddz. 225a potwierdza jego
przynale¿noœæ do Dryopteris cambriensis (Fraser-Jenk.) Beitel & W.R. Buck. Oba
taksony by³y dotychczas zaliczane do D. affinis agg.