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Folia Cryptog. Estonica, Fasc. 51: 61–66 (2014)
http://dx.doi.org/10.12697/fce.2014.51.06
INTRODUCTION
Due to increased forestry impact on biodiversity
worldwide, knowledge about species require-
ments for long-term existence in forest land-
scape is necessary. Epiphytic bryophytes and
lichens are good indicators to evaluate the forest
naturalness in Northern countries (Ikauniece et
al., 2012). Among these indicators are red-listed
species and ecology of these species are poorly
known. Forest fragmentation increases the dis-
tance between species populations affecting also
local microclimate (light intensity, humidity),
providing difculties for species survival (Bald-
win & Bradeld, 2007; Svoboda et al., 2010).
Therefore species dispersal ability (Jüriado et
al., 2011) and establishment conditions in a
new suitable patch are important drivers for
species distribution (Sillett et al., 2000; Snäll
et al., 2003). Transplantation experiments can
help to identify dispersal or microclimatic con-
ditions, which restrict the successful epiphyte
distribution between forest stands on suitable
substrates.
Transplantation experiments are needed to
predict factors inuencing the distribution of
epiphytic species, comparing the growth suc-
cess of model species between transplants in
Woodland Key Habitat (WKH) and in managed
forest. Lobaria pulmonaria (L.) Hoffm. and Nec-
kera pennata Hedw. are characteristic species of
old-growth nemoral forests (Lesica et al., 1991;
Ikauniece et al., 2012) and also suitable model
organisms for transplantation experiments (In-
gerpuu et al., 2007; Bidussi et al., 2013).
Gauslaa et al. (2001) conducted an experi-
ment with L. pulmonaria on wooden blocks, and
found that the growth of lichen was correlated
with rainfall during the studied time period and
L. pulmonaria was also susceptible to light after
transplantation. Edman et al. (2007) concluded
that selective cutting affected negatively the
abundance and frequency of L. pulmonaria fertil-
ity. Branches with transplanted L. pulmonaria
grew comparatively better in old-growth forest
with higher light than in managed forest (Cox-
son & Stevenson, 2007). In contradiction Hilmo
(2002) did not nd a difference in lichen Lobaria
scrobiculata growth after transplantation experi-
ments in old-growth forest and young planted
forest, conrming the hypothesis, that dispersal
limitation could be more important than micro-
climatic conditions in a particular forest stand
for this lichen species distribution. Hazell and
Gustafsson (1999) found, that highest survival
of L. pulmonaria was found in clustered trees on
the clear felled sites and the survival was similar
between the scattered trees and in clear-cuts
and in the forests. The vitality of L. pulmonaria
was highest on clustered trees in the clear-cut
and lowest in the forest (Hazell & Gustafsson,
1999). Antitrichia curtipendula (Hedw.) Brid. as
well as Neckera pennata are old-growth forest
bryophyte species. Hazell and Gustafsson (1999)
found that survival and vitality of transplanted
A. curtipendula was signicantly higher in forest
than in a clear-cut.
Different methods have been used for trans-
plantation experiments. Rosso et al. (2001) used
plastic net transplants in bags for biomass study
with A. curtipendula. Hazell and Gustafsson
(1999) used plastic nets with metal staples for
Transplantation experiments with Neckera pennata and
Lobaria pulmonaria in nemoral woodland key habitat
and managed forest
Anna Mežaka
Reasearch Institute for Regional Studies, Rezeknes Augstskola, Atbrīvošanas aleja 115, LV-4601, Rēzekne, Latvia.
E-mail: anna.mezaka@ru.lv
Abstract: Transplantation experiments with Neckera pennata and Lobaria pulmonaria were conducted in nemoral Woodland
Key Habitat and managed forest (in total 38 trees) in Latvia. Elastic cotton bandage was the best method for the present
transplantation experiments. Dispersal and establishment limitation were found to be crucial for N. pennata and L. pulmonaria
distribution between forest stands.
62 Folia Cryptog. Estonica
L. pulmonaria and A. curtipendula transplan-
tation. Gauslaa et al. (2006) used frames for
transplantation experiments with L. pulmonaria
on Picea abies (L.) Karst. in Norway. Ingerpuu
et al. (2007) made successful transplantation
experiment with Neckera pennata in Estonian
nemoral forests by pressing bryophyte into bark
crevices.
In boreo-nemoral zone bryophyte trans-
plantation experiments have rarely been done
and results of these studies may help to give
recommendations for nature protection and
forestry planning. The aims of the present study
are to nd the most appropriate transplanta-
tion method for studied nemoral forests and
to check if dispersal is more important for L.
pulmonaria and Neckera pennata distribution
than a change in forest management regime,
comparing nemoral WKH versus managed for-
est stand.
MATERIALS AND METHODS
Study area
Transplantation experiments were conducted
in Zīļu pļavas, Skrīveri district, Skrīveri village,
Mid Latvian Geobotanical region in nemoral
WKH (56o40'0''N 25o03'07''E) and in a nemoral
managed forest stand (56o40'14''N 25o03'07''E).
The WKH forest stand was 130 years old with
Fraxinus excelsior L. dominating, but recent cut-
tings had been made in managed forest stand,
where isolated F. excelsior in age between 50-100
years were left. Natural structures as coarse
woody debris in different decay stages, living
trees with different DBH were present in WKH.
N. pennata was growing also naturally in the se-
lected WKH. The distance between studied WKH
and managed forest stand was around 300 m.
Recently cut F. excelsior stumps were presented
in managed forest stand and only solitary F.
excelsior individuals were left. The territory in
transplantation experiment was selected based
on WKH inventory data.
Transplantation
Neckera pennata is autoicous epiphytic and
epilithic lowland moss. It is distributed in Cir-
cumpolar boreal-montane zone. This moss is
glossy yellowish green with undulate leaves,
and up to 10 cm long secondary shoots grow-
ing off the substrate with pH between 3.8–6.1.
The capsules are common, hidden by the leaves
(Smith, 2004; Strazdiņa et al., 2011).
Lobaria pulmonaria lichen thallus is pale brown
when dry and green when wet growing mainly
as an epiphyte. Lobes are 8–30 mm wide and
until 7 cm long with strongly ridged surface,
soralia are present in lobe margins and along
the thallus ridges. Isidia may be present among
soredia, apothecia infrequent. Good indicator of
rich, unpolluted, old forests (Brodo et al., 2001).
L. pulmonaria and N. pennata transplants were
taken from ve adjacent Fraxinus excelsior do-
nor trees with diameter at breast height (DBH)
of 1.00 m, located between the managed forest
and the studied WKH. All recipient F. excelsior
trees were selected randomly with similar DBH
(>0.23 m). Tree DBH varied among these trees
in WKH between 0.24–0.41 m and in the man-
aged forest between 0.29–0.40 m. Bark crevice
depth varied from 2.00 to 8.00 mm in WKH and
from 2.00 to 5.00 mm in the managed forest.
Tree bark pH varied between 4.59–6.21 in WKH
and between 4.08–5.63 in the managed forest.
Transplants of N. pennata and L. pulmonaria
were removed at 1.20 m height from the ve F.
excelsior donor trees. The length of N. pennata
transplants varied from 6.00 to 8.30 cm and
they were around 2 cm wide (2–3 secondary
shoots). The size of L. pulmonaria transplants
was around 9 cm2. All N. pennata transplants
were put at a height of 1.20 m on a tree in North
direction, and L. pulmonaria at a height 1.30 m
in North direction on receiving F. excelsior trees
in a managed forest and in the WKH. The lichens
and bryophytes growing on transplant contact
zone were removed before transplanting.
Transplantation was started in November
2006 with 10 transplants in managed forest
and 10 transplants in WKH. Transplants were
attached with shing line (tied two times around
the tree) for N. pennata and with combined
method (shing line and shing net) for L. pul-
monaria to recipient trees. One transplant unit
of L. pulmonaria and one of N. pennata were
transplanted to each tree. All transplantations
were conducted on the same day when they were
collected and each transplant was pulverized
with distilled water to decrease physiological
stress.
Checking of transplants was made in April
2007. Two receiving trees in the WKH and one
in the managed forest had fallen down. Also,
two lichen transplants in WKH and three lichen
transplants in the managed forest had fallen
63
down. New additional trees were selected instead
of dead trees and transplanted with the species.
In addition 10 new trees in the WKH and in
managed forest were selected in May 2007. Since
several transplants had fallen down, now the
other method, the cotton bandage was used for
attaching transplants. Cotton bandage consists
of cotton and elastic material used in medicine
and being available in pharmacy.
In total transplantation experiment was
made on 40 trees (20 in WKH and 20 in man-
aged forest).
The next observations were made in August
2007, December 2007, March 2008, August
2008, December 2008, and November 2009.
A digital photo was made of each transplant
at each inspection time. Transplants were pho-
tographed with a Powershot SX100 IS Canon
digital photo camera with 8.0 mega pixels,
Canon zoom lens 10xIS, and aperture 6.0-60.0
mm 1:2.8–4.3.
The vitality of transplants was evaluated
subjectively on a four grade scale based on
digital photographs: 1) high vitality; moist trans-
plant is greenish, without damaged patches or
margins, 2) medium vitality; transplant is still
greenish, but some damaged patches occur, 3)
low vitality; more than half of transplant area
is damaged, remnant green patches left, 4)
transplant is dead; transplant is brown, without
living tissues.
Data analysis
Transplantation data were analyzed with Chi-
square test in the R 2.7.2 package to test dif-
ferences between WKH and managed transplant
vitality. In total 19 Neckera pennata and 19
Lobaria pulmonaria transplants from managed
and from WKH were analyzed from April 2007
and November 2009.
RESULTS
Based on preliminary study in 2006, shing
line was not the best method for transplanta-
tion experiments since in total 5 transplants
from 20 were fallen down. Cotton bandage was
more appropriate transplantation method in
the present study – only six transplants from
40 were fallen down. Transplants xed with
cotton bandage were in the same position also
after three years. Transplants tied with shing
line had in several cases (especially transplants
of Lobaria pulmonaria) changed the angle or
location on tree.
In 2009 the highest percentage of Neckera
pennata transplants (52.63%) in the managed
forest belonged to second vitality class, but in
WKH to the third vitality class (63%; Fig. 1).
The highest percentage of Lobaria pulmonaria
transplants both in managed forest (44.44%)
and in WKH (63.16%) belonged to the second vi-
tality class. Completely dead transplants (fourth
Fig. 1. Transplants of Neckera pennata and Lobaria pulmonaria divided into vitality classes (1–3)
in managed forest and WKH at the end of the experiment.
64 Folia Cryptog. Estonica
vitality class) were found neither in managed
forest nor in WKH. The highest percentage of
transplants for rst vitality class was found in
L. pulmonaria (16.67%) in managed forest stand,
while N. pennata showed 15.79% of transplants
in rst vitality class in WKH.
No signicant differences were found in
Neckera pennata and Lobaria pulmonaria trans-
plants vitality between WKH and managed forest
(Table 1).
a managed and WKH forest stand. The present
study is in accordance with study by Öckinger et
al. (2005), where dispersal limitation with mean
dispersal distance 35 m was found for Lobaria
pulmonaria in deciduous forest of Southern
Sweden. The vegetative dispersal distance for L.
pulmonaria was found even shorter in Estonian
forests, 15–30 m between host trees (Jüriado
et al., 2011). The present study is in contradic-
tion with the study by Edman et al. (2007), who
found that selective cutting affected negatively
the abundance and frequency of Lobaria pulmo-
naria. L. pulmonaria was found to be sensitive
to light after transplantation experiments by
Gauslaa et al. (2001) and Gauslaa et al. (2006),
suggesting, that L. pulmonaria might die in open
habitats by high light during long dry periods.
Lobaria pulmonaria transplants grew compara-
tively better in old-growth forest with higher
light compared with managed forests (Coxson
& Stevenson, 2007). Other studies support the
results our study, e.g. limited dispersal was
found important for Lobaria oregana (Sillett et
al., 2000).
The present study shows, that dispersal and
establishment limitation are signicant vari-
ables shaping Lobaria pulmonaria and Neckera
pennata distribution in nemoral forest stands
since signicant differences were not nd in
transplant vitality between managed forest and
WKH. This should be taken into account plan-
ning nature protection and long term forestry
in landscape and forest stand scale. Retaining
potential substrate trees, especially Fraxinus
excelsior, for epiphyte distribution in nemoral
managed forests adjacent to nemoral old-growth
forests with species with limited dispersal,
transplantation and forest continuity could
ensure continuous existence of epiphyte spe-
cies, especially specialists with limited dispersal
abilities in boreo-nemoral zone.
ACKNOWLEDGEMENTS
The study was financed by projects ESF
2004/0001/VPD1/ESF/PIAA/04/NP/3.2.3.1/
0001/0001/0063, University of Latvia research
project ZP2008/ZP08, Latvian Academy of Sci-
ences grant 05.1512/147, Ministry of Educa-
tion, Youth and Sport of the Czech Republic,
Deutsche Bundesstiftung Umwelt Stipendium
für Nachwuchswissenschaftler. Thanks is given
to Guntis Brūmelis for comments in manuscript.
Table 1. Chi-square test between transplant
vitality classes in managed forest versus WKH.
Transplants x-squared df p
Neckera pennata vitality 5.33 3 0.15
Lobaria pulmonaria vitality 1.31 3 0.52
DISCUSSION
Fishing line and shing net are not exible
enough for fixing the transplants and also
not strong enough and are probably damaged
by winter frosts. During the experiment, the
shing line and shing net became slack after
one year (personal observation). Medical cot-
ton bandage was much more elastic and may
preserve its pressing property during harsh
winter frosts better. Based on the results of
the present study, the selected transplantation
method with a cotton bandage appears promis-
ing for transplantation experiments in the future
alongside with a plastic net with metal staples,
applied for Antitrichia curtipendula and Lobaria
pulmonaria transplantation in Sweden (Hazell,
Gustafsson; 1999).
Transplant vitality of Neckera pennata in
managed forest and WKH did not differ signi-
cantly in the present study. The present result
partly agrees with Ingerpuu et al. (2007), who
found that dispersal limitation is important
for Neckera pennata distribution, but not mi-
croclimatic conditions. Wiklund and Rydin
(2004) found that yearly precipitation was the
most important factor as well as establishment
properties for Neckera pennata colony growth,
but not forest site. Werth et al. (2006) hypoth-
esized that ecological conditions are crucial for
establishment of Lobaria pulmonaria instead of
dispersal limitation.
Signicant difference was not observed also
in vitality of Lobaria pulmonaria transplanted to
65
I am grateful to State Forest Service for help
with maps and access to database. Thanks are
given to the owners of the studied forest stands
in Skriveri district for understanding and pos-
sibility to do this research. Thanks are given to
anonymous reviewers, that helped to improve
the manuscript.
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