The structure of selected stands of Lycopodium annotinum L. on the

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The structure of selected stands
of Lycopodium annotinum L. on the grounds
of the Opoczno Forestry Management
ANNA ŒLIWIÑSKA-W YRZYCHOWSKA
Department of Ecology and Nature Conservation, Jan D³ugosz University
in Czêstochowa, Al. Armii Krajowej 13/15, PL-42-200 Czêstochowa, Poland;
e-mail: [email protected]
ABSTRACT : The aim of the study was to compare the structure of two stands of
Lycopodium annotinum occurring in fresh pine coniferous forest and fresh mixed
pine coniferous forest. Soil fertility, determined here as a habitat type of the forest,
is a factor significantly affecting the number and type of shoots produced by
Lycopodium annotinum. The statistical analysis proved that two patches of Lycopodium
occurring at different habitat can be treated as two separate coenopopulations
despite the small distance between them. In coenopopulations occurring in the mixed
coniferous forest there is a higher density of shoots with and without strobili comparing to the coenopopulation of fresh pine coniferous forest. Additionally, in the
latter coenopopulation the phenomenon of atrophy is more escalated and a higher
percent of shoots starts reproducing using spores.
ABSTRAKT : Celem badañ by³o porównanie struktury dwóch stanowisk Lycopodium
annotinum wystêpuj¹cych na siedlisku boru œwie¿ego i boru mieszanego œwie¿ego.
¯yznoœæ siedliska, okreœlona tutaj przez typ siedliskowy lasu, jest czynnikiem
istotnie wp³ywaj¹cym na liczbê i typ pêdów wytwarzanych przez wid³aka. Analiza
statystyczna udowodni³a, i¿ dwa wystêpuj¹ce na odmiennym siedlisku p³aty wid³aka,
mimo niewielkiej odleg³oœci pomiêdzy nimi, mog¹ byæ traktowane jako dwie odrêbne
cenopopulacje. W cenopopulacji rosn¹cej na siedlisku boru mieszanego wystêpuje
wiêksze zagêszczenie pêdów zarodnionoœnych i p³onnych w stosunku do cenopopulacji
z boru œwie¿ego. W cenopopulacji rosn¹cej na siedlisku boru œwie¿ego bardziej
nasilone jest zjawisko obumierania pêdów. W cenopopulacji tej wiêkszy procent
pêdów przystêpuje do rozmna¿ania przy u¿yciu zarodników.
KEY WORDS : Lycopodium annotinum, horizontal structure, vertical shoots, spatial
structure, modular growth
Œ LIWIÑSKA -W YRZYCHOWSKA A. 2008. The structure of selected stands of Lycopodium
annotinum L. on the grounds of the Opoczno Forestry Management. – In: E. Szczêœniak,
E. Gola (eds), Club mosses, horsetails and ferns in Poland – resources and protection. –
Institute of Plant Biology, University of Wroc³aw, Wroc³aw, p. 235–244.
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Anna-Œliwiñska-Wyrzychowska
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Introduction
Lycopodium species are worth of interest as descendants of formerly very
numerous Lycopodiophyta. The entire genus is under legal protection in Poland (Minister of Environment 2004). In the past, L. annotinum was collected
and used to make Easter decorations. The procedure is illegal now; however
in some parts of Poland it is still practised. At present, the area and the
number of stands of Lycopodium species lessen or even they completely
disappear because of some other reasons, too (Juchacz et al. 1993). These
reasons include the habitat changes, e.g. decreasing humidity of soils, and
economic use of forests (Piêkoœ-Mirkowa, Mirek 2003). The intensification
of cutting down the trees, which has been taking place for the last decade,
probably speeds up the process of disappearing of Lycopodium species stands
in manage forests. However it is difficult to determine unambiguously, which
factor is crucial. Thus, the continuous studies of the condition of these populations are necessary. The aim of the study was to compare the structure
of two Lycopodium annotinum stands of a similar size occurring in the habitat of fresh pine coniferous forest and fresh mixed coniferous forest.
1. Material and methods
The studies were conducted on the grounds of the Opoczno Forestry Management, Dêba Forestry. The approximate position of the study areas was:
51 o29’53” N, 20 o14’37” S. In Dêba Forestry, two neighbouring forest units
183c and 153i were chosen, where Lycopodium annotinum occurred. The
unit 183c is a habitat of fresh pine coniferous forest. The layer of trees is formed
by Pinus sylvestris with Quercus robur in the underwood. It is accompanied
by single specimens of Betula pendula and Alnus glutinosa. The undergrowth
is mainly covered by Vaccinum myrtillus (38.4% on average) followed by
Lycopodium annotinum (8.4%). The unit 153i is a habitat of the mixed pine
coniferous forest. The tree layer is dominated by Pinus sylvestris assisted
by Quercus robur. In the underwood there is Quercus robur and Frangula
alnus. Lycopodium annotinum affects the physiognomy of the undergrowth
significantly (36%) and is followed by Vaccinum myrtillus (16.9%).
Lycopodium annotinum is a clonal plant with crawling and forking shoots
forming polycormon. The length of individual branches can reach a few metres.
Short, forked, vertical branches grow out of long horizontal shoots. Strobili,
which are groups of sporophylls, are formed at the top of vertical shoots. The
enlarging of the area of individual stands is achieved by the growth of hori-
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237
zontal shoots of polycormon (Callaghan et al. 1986a,b; Grochowski 1956).
According to Sutherland and Stilman (1998), production of long horizontal shoots
increases the probability of placing a shoot in a suitable habitat for further
development.
In each of two forest units one study plot of the size of 200 square metres
was marked. The study plots were situated in such a way that they included
the biggest possible part of the club moss stand and its edge part. Each study
plot was divided into numbered squares using Greig-Smith grating (Faliñska
2002). Within each square different kinds of modules of Lycopodium annotinum
were counted, i.e. dead shoots and vertical living ones, and among the latter
shoots with and without strobili were distinguished. Every vertical branch without
strobili was counted as a shoot without strobili regardless of the number of its
further bifurcations. Every vertical branch was recognized as a shoot with strobili
if at least one strobilus was present. The results were statistically analysed
but only for squares in which Lycopodium occurred. The conformity of
Lycopodium distribution in samples with normal distribution was tested
with the use of the Shapiro-Wilk test (with p<0.01). For further analyses nonparametric tests were used because of the way of sampling and the fact that
studied data did not have normal distribution. The series test of Wald-Wolfowitz,
U Mann-Whitney test, normality Kolmogorov-Smirnov test and the test
of difference between two rates of structure (Stanisz 2006) were conducted
to verify the hypothesis that the two analysed samples come from different
populations.
2. Results
Living shoots
In both studied plots Lycopodium annotinum formed thick patches. Within
them the squares without Lycopodium were very rare (Fig. 1). The value
of the average number of living vertical shoots per 1 square metre was more
than three times higher for the patch of L. annotinum in mixed coniferous
forest (339.7 shoots/m2; Tab. 1) in comparison to the patch from the fresh forest
(105.1 shoots/m 2). The series test of Wald-Wolfowitz, U Mann-Whitney test,
and normality Kolmogorov-Smirnov test showed statistically significant differences (p<0.01) between the number of living vertical shoots of L. annotinum
occurring in 1 m2 in the mixed and fresh forests. In a 1 m 2 there were recorded
maximally 786 shoots in the mixed forest and 718 in the fresh forest. The stand
in the mixed forest was characterised by higher diversity of the number
of shoots occurring in 1 m 2 in comparison to the stand in the fresh forest
(Fig. 1).
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Without Lycopodium
0 living shoots/1m2
1 to 199
200 to 399
400 to 599
600 to 799
Fig.1. The amount of living vertical branches of Lycopodium annotinum at study plots (a
number of shoots per 1m2)
Ryc. 1. Zró¿nicowanie liczby ¿ywych pionowych pêdów L. annotinum na badanych
powierzchniach (liczba pêdów na 1m 2)
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239
Tab. 1. Characteristics of examined Lycopodium annotinum stands at fresh pine and mixed
pine coniferous forest habitats
Tab. 1. Charakterystyka badanych stanowisk Lycopodium annotinum w siedlisku boru
sosnowego œwie¿ego i mieszanego boru sosnowego
Data (Dane)
Number of plots with L. annotinum
(Liczba powierzchni z L. annotinum)
Number of plots with living shoots of L. annotinum
(Liczba powierzchni z ¿ywyni pêdami L. annotinum)
Total number of living shoot branches
(Ca³kowita liczba ¿ywych odga³êzieñ)
Average number of living shoots per one plot ± s.d.
(Œrednia liczba ¿ywych pêdów na powierzchni ± s.d.)
Maximum number of living shoots per one plot
(Maksymalna liczba ¿ywych pêdów na powierzchni)
Total number of shoots with strobili
(Ca³kowita liczba pêdów ze srobilami)
Average number of shoots with strobili per one plot ± s.d.
(Œrednia liczba pêdów ze srobilami na powierzchni ± s.d.)
Maximum number of shoots with strobili per one plot
(Maksymalna liczba pêdów ze srobilami na powierzchni)
Participation of shoots with strobili (%)
(Udzia³ pêdów ze srobilami)
Total number of dead shoots
(Ca³kowita liczba martwych pêdów)
Average number of dead shoots per one plot ± s.d.
(Œrednia liczba martwych pêdów na powierzchni ± s.d.)
Maximum number of dead shoots per one plot
(Maksymalna liczba martwych pêdów na powierzchni)
Participation of dead shoots (%)
(Udzia³ martwych pêdów)
Fresh pineforest Mixed pineforest
(Bór sosnowy
(Mieszany bór
œwie¿y)
sosnowy)
162
162
153
161
16 084
50 999
105.1 ± 95.3
316.8 ± 193.5
718
786
2498
3621
16.3 ± 21.8
22.5 ± 20.4
148
108
16.3
7.4
5260
4030
32.5 ± 22.2
24.9 ± 14.7
106
68
32.3
11.9
Shoots with strobili
In the study plot situated in the fresh coniferous forest there were 148 shoots
of L. annotinum with strobili (on average 16.3 shoots/m2; Tab. 1), whereas in
the mixed forest 108 of them (on average 22.5 shoots/m 2) were present. The
U Mann-Whitney and Kolmogorov-Smirnov tests showed statistically significant differences (p<0.01) between the number of shoots with strobili occurring in 1 m2 in the mixed and fresh forests. The most conservative Wald-Wolfowitz
test did not show statistically important differences between the two studied
stands. It suggests that despite very significant differences in the number of
living shoots, these two studied plots are not so different regarding the number of produced shoots with strobili. To compare the contribution of shoots
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240
Without Lycopodium
0 .00%
1 to 24%
25 to 49%
50 to 74%
75 to 99%
100%
Fig 2. Contribution of dead branches of Lycopodium annotinum in the total number of
branches at study plots
Ryc. 2. Udzia³ procentowy martwych pêdów L annotinum w ca³kowitej liczbie pêdów na
badanych powierzchniach
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The-structure-of-selected-stands-of-Lycopodium-annotinum
241
with strobili in both plots, the double-tailed test of differences between two
rates of structure showed statistically important differences with p=0.0149.
The average contribution of shoots with strobili in the plot in the fresh forest
(16.3%) was more than twice higher compared to the plot in the mixed forest
(7.4%). In the mixed forest stand Lycopodium invests much more in the growth
of shoots without strobili than in production of strobili in comparison to the stand
in the fresh forest.
Dead shoots
In the mixed forest, the significantly smaller percentage of dead shoots of
L. annotinum (11.9%) was observed than in the fresh forest (32.3% – Fig. 2,
Tab. 1). The double-tailed test of differences between two rates of structure
showed statistically important differences with p=0.000 in the contribution of
dead branches in both plots. Exactly the same situation was found when comparing the average number of dead shoots in 1 m2. U Mann-Whitney test showed
statistically important differences with p<0.01. Wald-Wolfowitz and KolmogorovSmirnov tests showed statistically important differences with p<0.03.
In both stands of Lycopodium, squares within which all observed shoots
were alive, very rarely occurred. There were also squares present in which
all shoots were dead. In both cases, squares were usually situated in the edge
parts of a Lycopodium patch (Fig. 2). It is worth mentioning that in the mixed
forest stand squares containing exclusively dead shoots, are grouped in the
“lower“ part of the picture (Fig. 2). Oppositely, squares with all Lycopodium
shoots living are grouped in the “upper“ part of the picture. It could indirectly
indicate the direction of Lycopodium expansion (“upper“ part of the picture)
and places, from which Lycopodium moves back (“lower“ part of the picture
– Fig. 2). In the mixed forest, the percentage of dead shoots in the studied
part of polycormons was only slightly changeable and apart from the edge squares
the remaining area seemed to be homogeneous in this respect (Fig. 2). These
relations are a little different in the study plot in the fresh coniferous forest.
Similarly to the mixed forest, squares in which there were only dead shoots of
Lycopodium were situated in the edge parts of polycormons. However, squares
with the high percentage of dead shoots were scattered at the whole area of
the stand.
3. Discussion
The mosaic picture, similar to that described above, occurred also in the
studied Lycopodium annotinum stands in fresh coniferous forests at Olkusz
Upland (Œliwiñska-Wyrzychowska 2008). In the edge parts of policormons young
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Anna-Œliwiñska-Wyrzychowska
242
shoots are produced, which are responsible for Lycopodium expansion (Cieszko
1999). If horizontal shoots find suitable conditions, they will root and fill
the space by producing side and vertical branches. In unfavourable conditions,
the horizontal shoot can grow until the transport of water and nutritious substances from the central parts of policormon is possible. Then the shoot dies
(Callaghan 1986a). Lycopodium annotinum is a modular species, which realizes guerrillas strategy of growth. By building up the number of modules within
each part of policormon, it can settle gaps appearing in vegetation (Callaghan
et al. 1986b). Such a way of expansion of L. annotinum occurs the most often
in coniferous forests (Juchacz et al. 1993; Œliwiñska-Wyrzychowska 2008;
Œliwiñska-Wyrzychowska, Bogdanowicz 2008; Œliwiñska-Wyrzychowska,
Ksi¹¿czyk 2008).
As previous studies in Czêstochowa and Olkusz Uplands showed, density
of various modules can differ significantly between different stands occurring at the same type of habitat (Œliwiñska-Wyrzychowska 2008; ŒliwiñskaWyrzychowska, Bogdanowicz 2008). Density of living shoots and shoots with
strobili was higher in stands occurring in the fresh coniferous forest habitat
compared to that of mixed forest (Œliwiñska-Wyrzychowska 2008; ŒliwiñskaWyrzychowska, Bogdanowicz 2008); this result is opposite to the one observed
in the present study.
Juchacz et al. (1993) noted in their stand a much bigger number of shoots
with strobili, which on average was 41,2 and 28,6 per m2 in two consecutive
years, than the one calculated in the present study. The mentioned above stand
was located between the edge of peat bog and a forest road at the edge of
mixed forest. The contribution of dead shoots noted in the present study in
both forest types was much higher than that observed in the stand at Czêstochowa
and Olkusz Uplands (Œliwiñska-Wyrzychowska, Bogdanowicz 2008; ŒliwiñskaWyrzychowska 2008). It can indicate the occurrence of the intensified process of shoot dying in both studied stands.
On the basis of this study results and the review of literature it is not possible to answer which type of habitats, the mixed forest or fresh pine forest,
is preferential for the presence of L. annotinum. L. annotinum is generally
noted in pine forests in the area of whole Poland but the number of its stands
is successively decreasing because of the intensified tree cutting. Because of
the small number of club moss stands we must consider the necessity of leaving fragments of forest stands in the areas of tree cutting with intact undergrowth. It may allow the club moss stand to survive and later, when suitable
conditions occur, to increase the possibility of club moss expansion and spore
dispersion to the neighbouring area.
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243
4. Conclusion
Fertility of the habitat determined here by the type of forest is a factor which
significantly affects the number and type of shoots formed by Lycopodium
annotinum. The statistical analysis proved that two patches of Lycopodium
occurring at different habitats can be treated as two separate coenopopulations
despite the small distance between them.
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Struktura wybranych stanowisk Lycopodium annotinum L.
na terenie Nadleœnictwa Opoczno
Celem badañ by³o porównanie struktury dwóch stanowisk Lycopodium
annotinum, wystêpuj¹cych na siedlisku boru œwie¿ego i boru mieszanego
œwie¿ego. Na stanowisku w borze mieszanym wystêpuje wiêksze zagêszczenie
pêdów zarodnionoœnych i p³onnych (na 1 m2) w stosunku do stanowiska z boru
œwie¿ego. Na stanowisku z boru œwie¿ego bardziej nasilone jest zjawisko
obumierania pêdów. Mimo mniejszej ca³kowitej liczby pêdów z k³osami
zarodnionoœnymi, na tym w³aœnie stanowisku wiêkszy odsetek pêdów przystêpuje
do rozmna¿ania przy u¿yciu zarodników. Analiza statystyczna (Test U MannaWhitneya, test Ko³mogorowa-Smirnowa przy p<0,01) wykaza³a istotne
statystycznie ró¿nice w liczbie ¿ywych pêdów pionowych, w liczbie pêdów
z k³osami zarodnionoœnymi oraz w liczbie pêdów martwych, wystêpuj¹cych
na porównywanych stanowiskach. Dla procentowego udzia³u pêdów z k³osami
zarodnionoœnymi oraz procentowego udzia³u pêdów martwych na obu badanych
stanowiskach, dwustronny test ró¿nic miêdzy dwoma wskaŸnikami struktury,
wykaza³ statystycznie istotne ró¿nice. Wyniki te pozwalaj¹ na wyci¹gniêcie
wniosku, i¿ dwa wystêpuj¹ce na odmiennym siedlisku p³aty wid³aka, mimo
niewielkiej odleg³oœci pomiêdzy nimi, mog¹ byæ traktowane jako dwie odrêbne
cenopopulacje. ¯yznoœæ siedliska okreœlona tutaj przez typ siedliskowy lasu
jest czynnikiem istotnie wp³ywaj¹cym na liczbê i typ wytwarzanych przez wid³aka
pêdów.