VEGETATION PATTERNS IN MANAGED AND SEMI-NATURAL BOREAL

FORESTS IN EASTERN FINLAND AND RUSSIAN KARELIA

Anneli Uotila

ACADEMIC DISSERTATION

To be presented, with permission of the Faculty of Forestry,

University of Joensuu, for public criticism in Auditorium H1 of Joensuu University,
Yliopistonkatu 2, on June 11th 2004, at 12 o’clock noon.

Faculty of Forestry
University of Joensuu
2004
 ISBN 952-458-470-0

Joensuun yliopistopaino 2004

 Abstract 3

ABSTRACT

Uotila, A. 2004. Vegetation patterns in managed and semi-natural boreal forests in eastern
Finland and Russian Karelia. Faculty of Forestry, University of Joensuu.

The impacts of tree populations or communities on the dynamics of understorey vegetation

are still poorly known, and the effects of trees and management in shaping the structure
of understorey vegetation are probably underestimated. In this context, this study aims at
investigating the structure of tree populations and understorey vegetation on mesic and
sub-xeric sites representing successional series of managed and unmanaged forests.
More specifically, the main objectives were: (i) identifying the main differences in stand
structure between managed and unmanaged forests; (ii) identifying the proper structural
features to indicate the degree of naturalness of tree stands; and (iii) analyzing how a
ground vegetation affected by management and timber harvesting may deviate from the
one that has developed under no management and timber harvest. The data for this
study represent the middle boreal zone, and it was collected in North Karelia, Finland, and
in Dvina Karelia, Russia, on both sides of the border between Finland and Russia.
The main findings of the study show that the “unmanaged” forests in eastern Finland
have signs of light selection fellings and slash-and-burn cultivation, and they are semi-
natural rather than natural. The degree of naturalness in the stand structure can be assessed
on the basis of cut stumps, diameter distribution of the tree stand and amount of coarse
woody debris. Fires and natural disturbances hold a key position in achieving a greater
resemblance to more natural stand and succession dynamics in managed forests, because
they maintain a continuity of coarse woody debris and uneven-aged stand structures.
Cuttings and other silvicultural treatments have decreased the abundance of liverwort
species in mesic sites and lichens in sub-xeric sites, and the coverage of dwarf shrubs
such as Calluna vulgaris in sub-xeric stands and Vaccinium myrtillus in mesic sites. At the
same time, the coverage of grasses has increased in both mesic and sub-xeric sites, and
the coverage of mosses in sub-xeric sites.

Key words: Boreal forests, ground vegetation, vegetation pattern, managed forests, semi-
natural forests

Author’s address: Faculty of Forestry, University of Joensuu, P.O.Box 111, FIN-80101

Joensuu, Finland, e-mail: [email protected]
4 Preface

PREFACE

The Finnish Cultural Foundation; the Foundation of North Karelia and the Foundation of
the University of Joensuu gave me the chance to initiate this study in spring 1994. The
Ministry of Agriculture and Forestry provided financial support for a period of one year.
The Faculty of Forestry, University of Joensuu, was mainly responsible for funding this
study, together with the Academy of Finland (Centre of Excellence Programme, project
no. 64308). The data on stand structure used in this study were partly collected in co-
operation with the project “Natural forests: structure, dynamics and biodiversity as
compared with managed forests”, funded by the Finnish Forest Research Institute. The
leader of this project, Mr. Antti Isomäki, encouraged me and provided valuable financial
and technical support. I wish to thank Harri Kontkanen, Juha Metros and especially Tapio
Ylimartimo for their co-operation and enjoyable times during the field work.
Metsähallitus assisted in the search for sample data from the Lieksa area. I also wish
to thank the North Karelian Border District. The border guards at the Inari and Ruuna
stations helped us with transportation and accommodation in the study areas located
along the border zone. I wish to thank Dr Raimo Heikkilä, the Research Centre of Friendship
Park, and Dr Sergei Tarkhov and Boris Kashevarov, the Kostamus State Nature Reserve,
for their help in the study areas in Dvina Karelia. Finally, but by no means the least, I
extend warm thanks to Mrs and Mr Santeri Lesonen in Dvina Karelia for their warm advice
and care during the Dvina excursions with my group.
I thank my supervisors Professor Seppo Kellomäki and Phil Lic. Juha-Pekka Hotanen
for their scientific support, advice and for their patience during the study. I wish to thank
the third supervisor professor Jari Kouki for his scientific advice. Part of the sample
collections was identified in the Botanical Museum of the University of Oulu and I wish to
thank them for their specialised know-how and pleasant co-operation. I would also like to
thank Ms Päivi Pulkkinen for some of the data calculations.
It has been a great pleasure to work among the innovative young people at the Faculty
of Forestry in Joensuu. I have shared my workdays, pain and joy with Maarit Similä and
Seppo Rouvinen for many years. I wish to thank Maarit and Seppo and many other
wonderful people for the fruitful discussions in the Borealis house and for their
encouragement to finish this work. I acknowledge John Derome, who revised the English
of this summary.
I am grateful to my friends for their continuous support throughout all my studies. My
colleague Tuula Smolander M.Sc. hiked and searched for sample plots in the Ruunaa
area, and comforted and encouraged me all these busy years. I wish to thank my family
and relatives, especially my wise parents. I thank Ilari Uotila, who encouraged me to
continue my studies and has taken care of our children during the field work.
Life is a gift, especially the new ones in nature. The best gifts in my life are my children
Heikki, Antti and Petteri. This thesis is dedicated to Heikki, Antti and especially to Petteri,
whose short life and life-work continues in this thesis. I thank my children for their main
scientific question “Why”, who have kept me in touch with real life and happiness.

Joensuu, February 2004

Anneli Uotila
 List of acticles 5

LIST OF ARTICLES

This thesis is based on the following articles, which are referred to in the text by the
Roman numerals:

I Uotila, A., Maltamo, M., Uuttera, J. and Isomäki, A. 2001. Stand structure in semi-
natural and managed forests in eastern Finland and Russian Karelia. Ecological
Bulletins 49: 149–158.

II Uotila, A., Kouki, J. Kontkanen, H. and Pulkkinen, P. 2002. Assessing the naturalness
of eastern Fennoscandian boreal forests. Forest Ecology and Management 161:
257-277.

III Uotila, A. and Kouki J. 2004. Understorey vegetation in spruce-dominated forests in

eastern Finland and Russian Karelia: successional patterns after natural and
anthropogenic disturbances. Manuscript submitted to Forest Ecology and
Management.

IV Uotila, A., Hotanen J.-P. and Kouki J. 2004. Succession of understorey vegetation in
managed and semi-natural Scots pine forests in eastern Finland and Russian Karelia
Manuscript submitted to Canadian Journal of Forest Reserch.

Mrs. Anneli Uotila is the principal author of all the articles in the thesis. She is also the
creator of the original ideas and planned the study sampling. In study I the data were
measured in co-operation with Mr. Antti Isomäki, the Finnish Forest Research Institute,
and Dr. Matti Maltamo carried out analyses. In study II, the data were collected by Mr.
Harri Kontkanen and Ms. Päivi Pulkkinen in co-operation with Mrs. Uotila. In studies II-IV,
Mrs. Uotila alone was responsible for the data analyses. She wrote the first drafts of the
articles, which were revised jointly by the co-authors.
6 Contents

CONTENTS

1. INTRODUCTION...................................................................................................... 7
1.1 Succession of vegetation ................................................................................ 7
1.2 Successional models ...................................................................................... 8
1.3 Secondary succession ....................................................................................9
1.4 Re-vegetation of vascular plants after disturbance ...................................... 10
1.5 Re-vegetation of bryophytes and lichens after disturbance ......................... 10
1.6 Aims of the study .......................................................................................... 12

2. MATERIAL AND METHODS ..................................................................................13

2.1 Study area and field sampling ....................................................................... 13
2.2 Historical databases...................................................................................... 15
2.3 Data analysis ................................................................................................. 15

3. RESULTS AND DISCUSSION ............................................................................... 18

3.1. Structure of tree populations in managed and semi-natural forests ............. 18
3.1.1 Variation of age and diameter distribution ........................................... 18
3.1.2 Tree recruitment ................................................................................... 19
3.1.3 Coarse woody debris ........................................................................... 19
3.2. Degree of naturalness ................................................................................... 21
3.3. Effect of management and successional stage on the
understorey vegetation ................................................................................. 22
3.3.1 DCA-ordination..................................................................................... 22
3.3.2 TWINSPAN-classification .....................................................................24
3.3.3 Summary of the DCA- and TWINSPAN-analysis..................................25
3.4. Patterns of understorey vegetation ...............................................................28
3.4.1 Species groups in general .................................................................... 28
3.4.2 Abundance of bryophytes .................................................................... 29
3.4.3 Abundance of lichens........................................................................... 31
3.4.4 Abundance of dwarf shrubs ................................................................. 32
3.4.5 Abundance of grasses ......................................................................... 33
3.5. Evaluation of results ............................................................................... 33

4. CONCLUSIONS .................................................................................................... 36

5. REFERENCES ....................................................................................................... 38

ARTICLES I-IV
 Introduction 7

1. INTRODUCTION

1.1. Succession of vegetation

The theory and descriptions of the forest site types applied in Finland are primarily based
on undisturbed mature forests (Cajander 1926, 1949) representing the climax stage in the
succession of plant communities. Nowadays, however, this kind of vegetation only rarely
exists or can be found (I). Forestry has modified the vegetation throughout the country,
with the consequence that most of the vegetation represents earlier phases of succession
(Finnish Statistical Yearbook of Forestry 2000). This results in problems in identifying the
site types, which could be avoided if the successional series behind the “climax” vegetation
would have been thoroughly investigated, as suggested by Kujala (1926b, 1979) and
Kuusipalo (1985). For example, the impacts of tree populations or communities on the
dynamics of understorey vegetation are poorly known, and the effects of trees and
management in shaping the structure of understorey vegetation are probably
underestimated (Kujala 1979, Kuusipalo 1985, Lahti and Väisänen 1987). The comparison
of succession series in managed and natural forests would allow one to identify how
management can modify the understorey vegetation (Cajander and Ilvesalo 1921, Cajander
1926, Pitkänen 1997, Pitkänen 1998), with consequent indications of the effects on the
diversity of the vegetation.
In the following, the succession of vegetation communities is defined as a change in
the species composition, structure or architecture of the vegetation over time (Wiegleb
1989). The succession of boreal forests is a combination of allogenic (controlled by external
factors) and autogenic succession (governed by biotic interactions within the community),
where openings in the canopy are created by the death of single trees or whole tree
stands as a result of external forces. Disturbances, either human-induced or natural,
maintain the successional dynamics in vegetation; in other words, natural disturbances
generate and maintain spatial heterogeneity in the understorey and tree layers (e.g. Sirén
1955, Ilvessalo 1967, Borman and Likens 1979, Oliver and Larson 1990, Linder 1997,
1998a, b, Chen and Bradshaw 1999, Kuuluvainen and Rouvinen 2000, Franklin et al.
2002, Karjalainen and Kuuluvainen 2002).
Watt (1947) applies the concepts of pattern and process in describing the dynamics
of vegetation. The concept of pattern refers to the structure of plant communities, and
the concept of process to the change in patterns between the time units of observations
as indicated with the help of successional stages, which are used to describe the overall
pattern of the succession. In this context, the concept of succession refers to the sequence
of changes in the vegetation that follows one another and results in a directional and
predictable change in the structure of the vegetation. Succession proceeds through primary
8 Uotila

succession, which starts from the situation where there are no plant communities, to
secondary succession, which represents the succession of vegetation replacing the primary
vegetation. Long-term changes in the vegetation at the landscape level are modified by
the climate (Van der Maarel 1988). This study deals only with secondary succession.
Disturbances that drive succession can be arranged in a hierarchy representing the
variations in magnitude and time scale (Pickett et al. 1987). Consequently, the succession
of plant communities represents variability of varying length. Over longer periods, the
succession of plant communities is often assumed to be cyclic; succession of the
vegetation returns again and again to the initial stage, which is followed by more mature
stages including the “climax stage” comprising different dominating species. Around this
trend, there are fluctuations in the structure of vegetation representing variations in
magnitude and time scale, e.g. fluctuation induced by gaps or pure random events, which
modify the overall pattern of the successional dynamics.
Disturbances like forest fires, wind- and snow-induced damages and clear-cuttings,
affect the secondary succession of vegetation. The dimensions of disturbance are space,
time and magnitude (Glenn-Levin and Van der Maarel 1992). The concept of space refers
to the extent, area, volume and location along the environmental gradient of disturbance.
In this context, the concept of time refers to the frequency and predictability of disturbance
(Reinikainen et al. 2000). The concept of magnitude can be measured in terms of the
force, i.e. intensity, or in terms of the degree of the effects, i.e. severity (White and Pickett
1985). In this study, it is assumed that fire in natural forests and clear-cutting in commercial
forests are the main disturbing factors controlling the dynamics of the ground cover. The
concepts of space, time and magnitude are applicable regardless of the disturbance
initiating the succession.

1.2 Successional models

Clements (1916) presented a theory of succession that states that succession is driven
through the phases 1) nudation, 2) migration, 3) ecesis, 4) competition, 5) reaction, and 6)
stabilisation, with the assumption that there is a climatic climax community at the end of
succession regardless of the other site factors. Gleason (1926) criticised strongly the
Clementsian theory, and emphasised the importance of the individualistic behaviour of
plant species and the role of random events in controlling the course of succession. In
contrast, Gleason (1926) claimed that no equilibrium assemblages occur in vegetation.
Odum (1969) and Margalef (1968) postulated that ecosystems have the tendency to develop
greater homeostasis, with the main emphasis on ecosystem energetics and the dynamics
of nutrient cycles.
 Introduction 9

The main view held since the early 1970s recognises that succession is primarily a
replacement process driven by the reactions of plants to plant-controlled (i.e. autogenic)
environmental modifications; succession proceeds in time along the gradients of resource
availability (Pickett 1976, Tilman 1985). This implies that vegetation change is the outcome
of fluctuating environmental conditions, and interactions between competitive species
with different life history characteristics (Pickett 1976, Noble and Slatyer 1980) or stochastic
processes (Horn 1974). This is well in line with the concepts of pattern and process used
by Watt (1947) in describing the succession of plant cover. He showed that plant
communities represent a mosaic of structural phases, which change from one phase to
another. The modern view of vegetation change emphasises that relatively frequent
disturbances are the main factors controlling the course of succession, and it accepts
that the course of succession is also controlled by stochastic events (Connell 1978, Huston
1979, Pickett and White 1985a).

1.3 Secondary succession

The secondary succession of boreal forests is often classified into two general phases
(Sirén 1955) with respect to tree populations or communities: the primary or first tree
generation (primary stand), and the second generation (secondary stand) that follows
primary generation. The primary stands develop after large-scale, catastrophic events
such as fire or clear-cutting. Secondary stands develop when the primary generation is
gradually replaced, in the absence of fire or other external disturbing factors, through
small-scale gap dynamics (Sirén 1955, Oliver and Larson 1990) that modify the properties
of the tree stand or patch. During the succession of primary and secondary stands, major
changes also concurrently occur in the ground cover in relation to the attainment of canopy
closure and stocking of trees.
Recently disturbed stands on mesic sites are typically characterised by pioneer species
that regenerate through wind-dispersed and buried seeds or through root sprouts (Kujala
1926a). These species are subsequently replaced by feather mosses and shade tolerant
vascular plants that primarily regenerate through root sprouts. In other words, large- and
small-scale disturbances create and maintain the diversity of microsites that affect the
number and growth of plants (Pickett and White 1985b, Crawley 1997, Økland 2000). In
general, the highest number of species is expected to occur at intermediate time spans
during secondary succession (Connell 1978, Huston 1979), and in the patches of
intermediate age with respect to the disturbance (Lahti and Väisänen 1987, Tonteri et al.
1990, Nieppola 1992, Tonteri 1994, Collins et al. 1995)
10 Uotila

1.4 Re-vegetation by vascular plants after disturbance

Vascular plants use a range of successional pathways, biological traits or tactics in

colonising a site after fire (Noble and Slatyer 1980, Longton 1992) and after clear-cutting
(Ross et al. 1986). However, the same pioneer and colonist species, e.g. Epilobium
angustifolium, Deschampsia flexuosa, Calluna vulgaris (Sarvas 1937, Ferm and Pohtila
1977, Nieppola 1992, Pitkänen 1998, Vanha-Majamaa 1998,), are usually present after
fire or clear-cutting. This implies that the vegetation prior to the disturbance on both burnt
and unmanaged and managed stands plays the key role in controlling the species
composition of the plant cover during succession after the disturbance (Halpern and
Spies 1995).
The pioneer plants or the dominants of the early succession regenerate from diaspore
or seed banks (Grime 1979, Clement and Touffet 1990, Behm 2001). On the other hand,
many species utilise the same dispersal mechanism as the seeds of Epilobium
angustifolium, and this species seems to be more effective in seed production and
regeneration after human-induced disturbances (Bråkenhielm and Liu 1998) than after
natural disturbances. Furthermore, the increasing availability of soluble nitrogen (e.g. NO3)
after clear-cutting also affects the vegetative growth of Epilobium angustifolium in the
same way as it effects on Deschampsia flexuosa (Foggo 1989, Högbom et al 2002). Similarly,
other environmental factors such as temperature and moisture (e.g. Uggla 1957, Viro
1969, Atlegrim and Sjöberg 1996, Pietikäinen and Fritze 1996, Jalonen and Vanha-Majamaa
2001) affect the regeneration and growth of vascular plants, resulting in differences in
succession between unmanaged and managed forests.
The resprouting response of some species is more vigorous to low-severity fire than
to high-severity fire (Schimmel and Granström 1996). These include aspen (Populus
tremula), goat willow (Salix caprea) and Deschampsia flexuosa, and dwarf shrubs such as
Calluna vulgaris, Empetrum nigrum and Arctostaphylos uva-ursi (Zackrisson 1977). On
the other hand, underground propagules are not easily destroyed by fire (Kujala 1926b),
e.g. Calluna vulgaris shows rapid regeneration from buried seeds after fire (Behm 2001).
In general, pioneer species make initially lifeless territories suitable for other species,
which implies that the suppression of fire in boreal forests has led to prolonged secondary
stands with increased dominance of climax species typical for the late succession
(Bradshaw and Zackrisson 1990).

1.5 Re-vegetation by bryophytes and lichens after disturbance

It is still poorly documented how the post-fire re-vegetation of cryptogams (particularly of

liverworts) deviates from that in clear-cutting areas, and what kind of environmental and
 Introduction 11

substrate cryptogams require in the different phases of succession (e.g. Foster 1985,
Longton 1992, 1997, Vellak and Paal 1999, Humphrey et al. 2002).
Bryophyte habitats are patchy in space and time, and the success of these species is
closely related to the prevailing properties of the habitats (Esseen et al. 1997, Söderström
and Herben 1997). In this context, the main divider between the managed and natural
forests is coarse wood debris, which is almost completely absent from managed forests
(Korhonen et al. 2001, Siitonen 2001, Rouvinen 2002). Coarse woody debris clearly
maintains the heterogeneity and functional diversity in forest ecosystems (Söderström
1988, 1993, Renvall 1995, Esseen et al. 1997, Martikainen 2000, Siitonen 2001, Rouvinen
2002), and this heterogeneity is almost completely lacking from managed forests
(Andersson and Hytteborn 1991, Kruyas et al 1999, Siitonen 2001).
Krusenstjerna (1945) divided bryophyte succession on decaying wood into the following
phases: liverworts are the first to appear, followed by competitive successful acrocarpous
mosses, and finally by pleurocarpous mosses. However, bryophyte life strategies (fugitive,
colonist and perennial stayer species (During 1992, Longton 1997)) also affect the
succession of the ground cover in both natural and managed forests. Unfortunately, only
a few studies have been carried out on how the life strategies differ, and on how cryptogams
modify the environmental conditions during succession after fire and clear-cutting (e.g.
Longton 1992, Mäkipää 2000b, Rees and Juday 2002).
The colonisation strategies of bryophyta are mainly based on regeneration through
the diaspore bank (Jonsson 1993a, During 1997) or in the case of lichens from asexual
soredia and the fragmenting thalli of lichens (Topham 1977). Short-lived fugitive species
like Ceratodon purpureus and Funaria hygrometrica become established during the first
year after burning (Foster 1985). In naturally uprooted patches, species such as Pohlia
nutans may occupy the site early in succession and remain among the forest floor
bryophytes for more than 100 years (Jonsson and Esseen 1990, Jonsson 1993a). Dominant
bryophyte species, which usually dominate in late succession, such as Pleurozium
schreberi and Dicranum sp., survive clear-cutting more successfully than fire (Jalonen
and Vanha-Majamaa 2001, Rees and Juday 2002).
The competition capacity of lichens is usually low (Kershaw 1977, 1985, Topman 1977),
and lichens mainly occupy dry sites where there is an absence of competition from other
species. Low water availability, high calcium content and low nutrient availability (Topham
1977), may favour lichen-rich vegetation. This means that the conditions on a site after a
fire and after clear-cutting are relatively different for lichens (Mälkönen and Levula 1996).
Since shading by trees easily suppresses lichens (Topham 1977), the role of ground lichens
in the plant cover probably diminishes during the gradual attainment of canopy closure
and stocking of trees, which occur in the development of a tree stand (Ahti 1959, Kershaw
1977).
12 Uotila

1.6. Aims of the study

The impacts of tree populations or communities on the dynamics of understorey vegetation

are still poorly known, and the effects of trees and management in shaping the structure
of understorey vegetation are probably underestimated. In this context, this study aims at

(i) identifying the main differences in stand structure between managed and unmanaged
forests;

(ii) identifying the proper structural features to indicate the degree of naturalness of
tree stands; and

(iii) analyzing how a ground vegetation affected by management and timber harvesting
may deviate from the one that has developed under no management and timber
harvest.

The study concerns only plants, although many issues in succession are connected to
different animal groups. Fungal and other pathogens may effect on the succession, but
these effects have been excluded. The material represents mesic sites of the Myrtillus
type dominated by Norway spruce (Picea abies (L.) Karst.) and sub-xeric upland forests
(dryish heaths) dominated by Scots pine (Pinus sylvestris L.).
The concept of natural forest refers in this study to forest vegetation established after
fire with no effect of management or harvesting. Truly natural forests are practically non-
existent in the region concerned, but early impacts were characterised by the removal of
single trees in light selection fellings (Lehtonen 1998, Aarnio 1999, Pitkänen 1999, Uotila
et al. 2002). Thus, the concept of semi-natural forests is preferred when considering forests
lying outside the scope of regular management and timber harvesting. The concept of
managed forest refers to forest vegetation affected by forestry, with cuttings and
management representing the contemporary patterns. This implies that the mature
managed stands included of this study are only lightly cut selectively, whereas younger
stands have experienced soil preparation, planting and other management measures after
clear-cutting. The material for the present study was collected in Northern Karelia (Finland)
and in the Dvina region of Karelia (Russia), both regions that represent the middle boreal
forest vegetation zone.
 Material and methods 13

2. MATERIAL AND METHODS

2.1 Study area and field sampling

The data used in this study include 38 sub-xeric stands dominated by Scots pine (Pinus
sylvestris), and 39 mesic upland soils stands dominated by Norway spruce (Picea abies).
The stands were measured during the summer in 1994-1998. The stands are located in
eastern Finland and in the neighbouring Karelian Republic, Russia (Fig.1), both areas

Study areas

Fig.1. Location of the study areas in eastern Finland and Russian Karelia. Vegetation
zones are according to Kalela (1961) and Ahti et al. (1968).
14 Uotila

representing the transition between the northern and middle boreal type of vegetation
(Kalela 1961, Ahti et al.1968). Consequently, the Vaccinium vitis-idaea forest site type
group (sub-xeric/dryish heath) is characterised by the features of both the Vaccinium type
and Empetrum- Vaccinium type (Cajander 1949, Keltikangas 1959). Similarly, mesic forests
are the Vaccinium myrtillus site type group, and they characterised by Myrtillus and
Vaccinium-Myrtillus types (Cajander 1949, Keltikangas 1959) dominated by Vaccinium
myrtillus.
This study utilises the temporary sample plots used to compile the chronosequences
representing successional series used in the study. The sample plots were located in sites
or stands not protected (managed) or protected (unmanaged). Four series were complied:
one for managed and unmanaged stands on the sub-xeric and one for managed and
unmanaged on the mesic sites, each series covering the whole succession series from
open areas to full maturity. For analytical purposes, the series were divided into five
successional stages: 1 = open/burnt, 2 = sapling, 3 = young, 4 = mature, 5 = over-mature/
old-growth regardless of management (I, II, III, IV). The classification was based on the
dominant height (mean height of the 100 tallest trees per hectare) and age of tree population.
In compiling the successional series, the managed stands were no problem compared
to the unmanaged stands. Especially, the young stands with no form of management
were in short supply in eastern Finland. This is why the unmanaged sapling and young
stands are partly located in the Kostomuksha Nature Reserve and the Kalevala National
Park, both in Russian Karelia (I, II, III, IV). On the other hand, the unmanaged stands on
the burnt sub-xeric sites were thinned one year before the fire in 1992. In other respects,
the unmanaged stands are well representative for succession under the natural conditions.
These stands are characterised by fire stumps, several storeys of trees with fire marks on
their bark, and large amounts of coarse woody debris and only a few signs of silviculture,
i.e. more than 60 years have elapsed since any cuttings or management (II).
The measurement of stand and sapling variables was based on fixed-radius, circular
sample plots. The size of the plots varied from 900 m2 to 2500 m2, depending on the
number of standing stems per plot (I). Tree species and diameter at breast height (dbh at
1.3 m) of each living (dbh>3 cm) and dead (dbh>10cm) tree were measured. Height and
age of the sampled living trees were determined, and the number of cut and natural
stumps on the sample plots counted (I).The number of seedlings with a minimum height
of 10 cm to saplings with a maximum height of 300 cm (dbh <30 mm) was counted on
sapling sample plots with a size of 100 m2 (Fig. 2, III, IV). Each sapling sample plot was
divided into 20 sectors, each with the same area (Isomäki 1995). The height of the tallest
sapling of each species was measured in every sector of the sapling sample plots. The
number of sapling sample plots varied from 3 in managed to 5 in unmanaged forests.
The understorey vegetation (available from 77 stands) was sampled in 10 quadrats
(each 1 m2) located on fixed-radius, circular sample plots. The quadrats were located
 Material and methods 15

systematically in the direction of the main and half-cardinal points (III, IV). The abundance
of each species of vascular plant, bryophyte and ground lichen was recorded in terms of
coverage (%) and frequency (III, IV). All species were determined. The nomenclature follows
Hämet-Ahti et al. (1998) for vascular plants, Vitikainen et al. (1997) for lichens and Koponen
et al. (1977, 1998) for mosses.

2.2 Historical databases

The history of the study areas was determined from maps drawn up for the enclosure of
common forest (1:8000) dating from the period 1844–99 and for forest land (1:20000), and
the records associated with these maps dated after 1909. The location of the field plots
on the old forestry maps was determined by checking the previously assigned forest site
type and stand structure. The archival catalogue of stands marked for cuttings and cutting
reports indicated the amount and type of cuttings on each plot (II). These results were
compared with the number of stumps and fallen and decaying trees. The location of one
third of the sample plots was identified on the land enclosure maps, the maps showing
whether the area had been under management. Information about slash-and-burn
cultivation, classification into the categories of a forest or a field, artificial regeneration,
soil preparation, removal of saplings, silvicultural treatments, and records of pruning,
fertilisation, work plans, annual reports and reports of forest fires were obtained from the
forest archives (II).
A dendroecological method was also used to identify the year of fires on the available
sample plots (Lehtonen 1997, 1998, Lehtonen and Kolström 2000). Interpretation of the
management history was also made with the aid of information about local (Potinkara
1993, 1997, Aarnio 1999, Pitkänen and Huttunen 1999) and national (Ahola 1959, Leikola
1987a, 1987b) forest history. Historical information was compared with the measured
variables. The final interpretation of the type of management was made after all the data
sources had been consulted. In the final phase the degree of certainty that the treatments
had actually been carried out was classified on the basis of the available records as
follows: the treatments were verified as having been carried out, the treatments were
possible verified, and the treatments had not taken (II).

2.3 Data analysis

Statistical significance of the differences between the treatments and successional stages
and their interaction terms (management successional stage) was tested by two-way
factorial ANOVA models (analysis of variance) using the stand structure and vegetation
16 Uotila

variables (I; III, IV, SAS User's Guide 1990, Anon. 1999). The mean value of the cover
percentages recorded in the ten quadrats of each circular field sample plot was used in
presenting the results and analysing the vegetation data. The vegetation was divided into
seven classes according to their life forms as follows: 1. Dwarf shrubs including the family
Lycopodiaceae (Kujala 1979) and dwarf-like Orthilia secunda, 2. Grasses including the
families Poaceae, Cyperaceae and genera Luzula, 3. Herbs, 4. Tree and shrub species, 5.
Ground lichens, 6. Mosses, and 7. Liverworts. The structural differences between the
semi-natural and managed stands in the same successional stage were also analysed
with the help of the post hoc tests of Tukey's HSD (III, IV, Anon. 1999).
Detrended correspondence analysis (DCA) was used to explore the structure of species
assemblages on the study areas in order to illustrate the combined effects of treatments
and successional stages (III, IV). DCA is a modification of correspondense analysis; i.e. a
metric scaling method based on a unimodal statistical model for the relationship between
species abundance and underlaying gradients (Økland 1990). DCA uses a chi-squared
distance measure with rescaled axes and downweighting rare species. The PC-ORD
program (McCune and Mefford 1999) has corrected the criteria of stability. A criterion of
tolerance is 0.0000001, and the maximum number of iterations is 999.
DCA ordinates both the species and samples simultaneously. This facilitates the
analysis on how the ground vegetation is affected by the management and timber harvest
may deviate from the one that has developed under no management and timber harvest.
The available data is suitable for this analysis, since the length of first axis varied from 2.9
to 4,3 S.D. units in the data of mesic and sub-xeric sites and in the combined data over
the sites. Furthermore, the Principal component analysis (PCA) was used in the analysis,
but it failed to group the vegetation in a proper way. PCA assumes that the species
abundances can be described a linear function (Økland 1990), but this was not the case
in the present data.
The analysis of DCA was further elaborated with the help of TWINSPAN classification
in order to identity in details and the differences between the semi-natural and managed
stands and to identify the plant species with value for indication of plant species occurring
in the similar conditions as regards the site type, management and maturity of the tree
stand (Gauch 1982). As regards the first axis TWINSPAN divides the ordination space into
two groups of plots, which helps to identify the plants or plant groups characteristic to the
succession ground cover under management or under no management. This is of primary
importance when assessing how the succession in the managed forests is deviating from
that in the natural forests as regards the present data.
TWINSPAN is based on dividing up a reciprocal averaging ordination space. The
vegetation data were classified by two-way indicator species analysis, which is a
hierarchical clustering method that uses differential species to characterise and separate
the classes (McCune and Mefford 1999). The accepted parameter values were: (i) a
 Material and methods 17

minimum group size of five plots for division, (ii) five indicator species, (iii) six levels of
division, and cut levels of octave values, i.e. 0 %, 0.5 %, 1 %, 2 %, 4 %, 8.0 %, 16 %, 32
% and 64 % for pseudospecies. For further details of the applications of TWINSPAN
classification, see Gauch and Whittaker (1981).
18 Uotila

3. RESULTS AND DISCUSSION

3.1. Structure of tree populations in managed and semi-natural forests

3.1.1 Variation in the age and diameter distribution

In the semi-natural stands, the age distribution of the trees was wider than that in the
managed stands regardless of site fertility (I, II). This was especially the case in the young
stands on the Russian side of the border (I, II, see also Syrjänen and Kalliola 1994,
Kvavchenko et al. 2000). In the semi-natural stands, the maximum number of age classes
was three (II), but in the managed stands the number of age classes was more than one
only in the most mature stands (II). The maximum age of the trees was the highest in the
burnt stands, indicating that some of the trees may have survived the fire and thus
represented the pre-fire tree stock.
The wide age distribution implies a broad range in the number of tree storeys and in
the diameter distributions. However, there were more tree storeys in old, managed stands
than in semi-natural stands. This implies that thinnings have modified the diameter
distribution by producing gaps in the stands, thus allowing the development of understorey
tree layers (see also Bailey 1996, Angelstam and Petterson 1997), i.e. the gap-forming
effect of thinning seems to be more effective than that of natural gap dynamics. In thinning
operations, the soil surface may also be disturbed and provide more favourable site
conditions for seedling establishment than in the case of natural gap formation (cf. Pohtila
1977, Angelstam and Petterson 1997).
In the semi-natural young stands, the diameter of the trees varied within a relatively
high range (II), but the total number of trees was higher in the managed sapling stands
than in the unmanaged ones (II). The diameter distributions in the managed sapling and
young stands were unimodal, but in the semi-natural stands the distributions were broader
with a smaller peak for larger diameters (I). The role of smaller diameter classes is dominating
in the distribution, as is typical for natural forests (I, II). The peak for the larger diameter
classes indicates that the older tree generations survived the fire (Sirén 1955, Franklin et
al. 2002, Karjalainen and Kuuluvainen 2002). This pattern was especially clear in the young
semi-natural stands (I), and it was recognisable also in the mature stands but not as
clearly as in the young stands. The dominance of small diameter classes in the distribution
indicates the absence of management during most of the life time of the stands. This
distributions pattern was more obvious in mesic forests than in sub-xeric sites (I (Figure
2), II (Figure 4)). However, the old-growth stands on the Finnish side of the border were
only 150-200 years old, but they still developed rather naturally following slash-and-burn
cultivation (Lehtonen 1997, Aarnio 1999). This is the case also in Dvina Karelia (Bazegskij
1998, Gromtsev 1998, Lehtonen and Kolström 2000, Karjalainen and Kuuluvainen 2002).
 Results and Discussion 19

In both areas, wild fires, natural or related to human activities, have also affected the
development of forests (Lampainen et al. 2002), with a consequent increase in the variability
of the stand structure (Wallenius et al. 2002). There is a weak positive correlation between
the occurrence of stumps and the volume of coarse woody debris in the Viena wilderness
area (Karjalainen and Kuuluvainen 2002), indicating that human activities have affected
the growth and development of these forests, but has not fully curbed the formation of
coarse wood debris.

3.1.2 Tree recruitment

In the mesic sites, the total number of seedlings and saplings was larger in the unmanaged
young stands (46 000 seedlings ha-1; height 10-300 cm) than in the managed ones (13 500
seedlings ha-1, Fig. 2 and see also III). This was also true for the sub-xeric sites dominated
by Scots pine. In the mesic sites, the density of seedlings was of approximately the same
magnitude (18 400 seedlings ha-1; height 20-200 cm) as reported by Lampainen et al.
(2002) for the Viena wilderness area. Irrespective of the site fertility, the recruitment dynamics
was dependent on the management, as indicated by the tree species composition and the
number or abundance of different species in the young stands (Fig. 2, III, IV).
In the mesic managed sites, deciduous seedlings competed strongly with coniferous
seedlings (Fig.2), suggesting that management has enhanced the recruitment and growth
of deciduous trees (Fig. 2). Especially, the occurrence of rowan (Sorbus aucuparia) was
high: from 42 to 58 % of the total number of saplings (Fig. 2). In contrast, the proportion of
coniferous trees in the mesic semi-natural stands was high (36 %), most of which were
Norway spruce (Picea abies). In the sub-xeric semi-natural saplings experienced crown
fires the proportion of Scots pine (Pinus sylvestris) seedlings was more than one half
(66%) of the total number of seedlings. Aspen (Populus tremula) occurred frequently in
the sapling stands (20 % of the total number of seedlings and saplings) on dryish heaths
and burnt mesic sites (60 %), but the proportion of aspen was relatively low in the other
semi-natural stands.

3.1.3 Coarse woody debris

Regardless of the site fertility and the phase of succession, the average volume of coarse
wood debris was greater in the semi-natural forests than in the managed ones, and it
increased along with increasing age of the stand with the maximum immediately after a
fire in a mature stand (Fig. 3, I). The result is comparable to that reported by Fridman and
Walheim (2000), Siitonen (2001), Korhonen et al. (2001), Sippola et. al. (1998, 2001), Pedlar
20 Uotila

et al. (2002), and Rouvinen et al. (2002a, b). The fire intensity was probably directly reflected
in the amount of coarse wood debris with the effect of the tree regeneration. Hytteborn et
al. (1987), and Hofgaard (1993b) have, for example, demonstrated that the presence of
decaying wood enhances the regeneration of trees in mesic boreal forests through providing
favourable sites for seeds to germinate and for seedlings to grow. Fires enhance the
natural mortality of trees with a pulse of dead wood in each fire episode. This is of primary
importance for supplying dead wood to maintain the presence of coarse woody debris

Fig. 2. The number of saplings ha-1 (height, 10- 300 cm) on mesic (a, b) and sub-xeric (c,
d) sites in managed (a, c) and semi-natural (b, d) stands of five successional stages (I, II,
III, IV, V). I= open/burnt, II = sapling, III = young, IV = mature, V = over-mature/old-growth.
 Results and Discussion 21

and to provide conditions favourable for specialised species or species with strict habitat
requirements (I, III; IV, Gustafsson and Hallingbäck 1988, Söderström 1988, 1995,
Andersson and Hytteborn 1991, Hansen et al 1991, Longton 1992, Esseen et al. 1997,
Jonsell et al. 1998, Kuusinen et al. 1998, Kruys 1999, Martikainen 2000, Kouki et al. 2001,
Siitonen 2001, Franklin et. al. 2002, Humphrey et al. 2002, Similä et al. 2002, 2003).

3.2 Degree of naturalness

The results strongly suggest that natural stand dynamics has not been the only factor
modifying the forests in the Finnish nature reserves and national parks. The unmanaged
forests of nature reserves and national parks in eastern Finland can be defined as any
forest that have not been subjected to silvicultural treatments for the past 50 - 100 years
(II). Fellings were verified on 33 % of the unmanaged sample plots, 21 % of the plots had
possibly been managed, and the forests in 46 % of the cases were completely untouched.
Mesic, old-growth stands were less affected (73 % completely untouched), but the tree

Fig. 3. The volume of coarse woody debris (CWD) and standing and downed trees in m3/
ha (a, b) and living trees, m3/ha (c, d) in mesic (a, c) and sub-xeric (b, d) sites in managed
(M) and semi-natural (SN) stands of five successional stages (I, II, III, IV, V, see Fig. 2).
22 Uotila

volume data, number of tree storeys and signs of slash-and-burn cultivation represented
clear evidence of the impact of human activities (II). On the mesic and sub-xeric sites, the
old-growth forests were uneven-aged (50-200 years old) with two stories, even though
these forests have been subjected to light selection fellings (II).
In general, the forests in North Karelia are, after the cessation of slash-and-burn
cultivation, semi-natural rather than natural ones. On the Russian side of the border, the
human impact is smaller as indicated by the higher number of tree stories. In the Vienansalo
wilderness area, on the other hand, the forests have been subjected to selective cuttings
(e.g. Yakovlev et al. 2000, Lehtonen and Kolström 2001, Karjalainen and Kuuluvainen
2002, Lampainen et al. 2002, Rouvinen et al. 2002b), as indicated by the stumps on the
sample plots. The mean number of cut stumps varied from 9 stumps ha-1 (Lampainen et
al. 2002) to 14 stumps ha–1 (Karjalainen and Kuuluvainen 2002), and to 26 stumps ha-1
(Rouvinen et al. 2002b) in the study area of Vienansalo. This is in line with the documents
concerning the management history of the area (I, II). Even the most natural areas are
more or less affected by man, who has utilised these areas for different purposes for
centuries. Stumps took more than 100 years to decompose in the middle boreal zone of
Norway (Storaunet et al. 2000, Groven et al. 2002). Logs decomposed completely within
70 years in the middle part of Sweden (Hytteborn and Packham 1987), but logs were still
visible after 150 years at higher latitudes (Dynesius and Jonsson 1991, Hofgaard 1993a).
In this context, the number of cut stumps and amount of woody debris represent meaningful
quantitative tools for identifying the degree of naturalness for practical forestry and
conservation (II), as demonstrated previously by Rouvinen et al. (2002b).

3.3. Effect of management and successional stage on the understorey vegetation

3.3.1 DCA ordination

In the ordinations, burnt dryish heaths and burnt mesic stands were located in the top
right corner (Fig. 4), and dryish semi-natural sapling stands on the left-hand side. In this
group, most of the sample plots in Russia were located on the left, which reflected the
differences in the abundance of lichens between the vegetation in burnt sapling stands in
Finland and in Russia (IV).The semi-natural sapling stands in the Ruunaa Conservation
Area in Finland have, however, been selectively logged after a crown fire that occurred in
1946 (II), while in Russia the dead trees are still standing. The volume of living trees,
occurrence of Norway spruce and the dominant height of the trees correlated with stand
age. The first axis reflected mainly the successional gradient and perhaps is connected to
the severity of disturbance. The increasing severity seems to increase light but moisture,
the basal area and the height of dominant trees seem to decrease along the second axis.
 BA basal area
BAdec basal area of deciduous trees
BAsp basal area of Norway spruce
CWDstand the volume of standing coarse woody debris
Hdom height of dominant trees
Vliv volume of living trees
Vsp volume of living Norway spruce
Vdec volume of deciduous trees
Vbi volume of living birch

Stage Managed Semi-natural

mesic sub-xeric

I ! "
II ! "
III ! "
V ! "
V ! "

Results and Discussion 23

Fig. 4. Ordination diagram of DCA with mesic and sub-xeric sites. Eigenvalues for the first and second axis are 0.468 and 0.115. The
total inertia of ordination is 2.2825. The direction of the vector shows the gradient in the environmental variable across the ordination
pattern and its length the strength of correlation. The symbols for all the stands decrease in size from stage I to stage V. Circles
denote managed stands and squares semi-natural stands. Open symbols are mesic stands and dark symbols are sub-xeric stands.
24 Uotila

Furthermore, the older mesic sample plots were positively correlated with the volume of
deciduous trees. Managed mature and overmature mesic sample plots are relatively close
to each other. The same result was found in article I and in the historical records (II), and
they are in accordance with the results of Rouvinen (2002). Old, selectively logged stands
resemble near natural stands (III, IV). Thus, the naturalness of forests is a continuity (see
also Haila 1995, Peterken 1996, and Rouvinen 2002), which represents the wide variability
of human influence on forest development.

3.3.2 TWINSPAN classification

The TWINSPAN classification produced a cluster hierarchy with six levels (Fig. 5). In the
dendrogram, the indicator species for each division are shown in decreasing order of
significance. At the 1st level, the sample plots were divided into the mesic (number of
plots in the cluster N = 45) and sub-xeric sites (N = 31), with Deschampsia flexuosa as an
indicator for the mesic sites and Calluna vulgaris, Cladina arbuscula and Cl. rangiferina for
the sub-xeric sites.
At the 2nd level, the left-hand subset represented the burnt stands in the Kitsi area
(N = 5), and the mesic sites elsewhere (N = 40). The indicator for this division was Dicranum
scoparium, with Deschampsia flexuosa, Dicranum polysetum, Hylocomium splendens (cut
level 1 %) and Pleurozium schreberi (cut level 4 %). The right-hand subset was
characterised by Blepharostoma trichophyllum, a liverwort growing on decaying wood in
moist and shady sites (Piippo 1993). The subset represented the area affected by a crown
fire in the Kostomuksha Nature Reserve (N = 2). The differential species were colonising
species like Calluna vulgaris (cut level 32 %), Populus tremula, Polytrichum juniperinum,
Pohlia nutans (cut level 4 %), and many Cladonia (e.g. Cl. clorophea, Cl. cornuta,
Cl. deformis, Cl. coccifera) species. The other plots (N = 29) represented stands of various
age in the sub-xeric sites, irrespective of management.
The left-hand side of the 3rd level (N = 5) represented the burnt semi-natural mesic and
sub-xeric stands, as indicated by Bryum capillare. Negative preferential species in the
first cluster were Vaccinium vitis-idaea (cut level 2 %) and the pioneer species Polytrichum
juniperinum (cut level 4 %). The next division of the 3rd level was indicated by Vaccinium
myrtillus, and its left-hand group mainly includes semi-natural older mesic stands
(N = 31). The right-hand side of this division mainly consisted of clear-cuttings (N = 9)
characterised by Polytricum juniperinum and Epilobium angustifolium representing the
pioneer species (N = 8). Common species in sub-xeric stands, Dicranum scoparium,
D. polysetum and Pleurozium schreberi, were positive differential species (N = 21).
At the 4th level (N = 5), Vaccinium uliginosum was the indicator of the first cluster, with
young semi-natural stands in the Viena study area. Vaccinium uliginosum may indicate
 Results and Discussion 25

the occurrence of paludification after fire. On the right-hand side, the coverage of lichens
like Cladina arbuscula indicated natural dryish heaths in both the second cluster (N = 4,
plots located in Russia) and the fourth cluster (N = 16). In the latter case, Cladina arbuscula
and Cetraria islandica were indicators of the semi-natural sample plots in Finland. In the
third cluster, Deschampsia flexuosa, Vaccinium uliginosum and Hylocomium splendens
were indicators for the division.
At the 5th level, Bryum sp. indicated clusters that may belong to the same mesic
group. In the second cluster, Brachythecium sp. indicated the presence of litter (Bates
2000, Söderström 1988) of deciduous species like Betula sp. They also indicated the
presence of dead wood . Melampyrun pratense distinguished the next clusters (N = 3),
mainly consisting of young (stages I and II) managed stands in the mesic sites. On the
right-hand side, the dryish heaths were a mixture of mature and over-mature stands,
either managed or semi-natural. Furthermore, these stands have experienced selection
logging, and they are “open” with plenty of light (II).
The 6th level represented the mesic cluster (N = 16), where semi-natural and managed
old sample plots are located in the same group. Hylocomium splendens was an approximate
indicator of a high degree of naturalness. The managed stands in this cluster have not
been managed for 20-30 years (II), and subsequently have a relatively large amount of
decaying wood (Fig. 3, I). These stands have clearly been subjected to management, as
indicated by the presence of cut stumps (II) and lower stocking (I). Similarly, the cluster of
sub-xeric stands consisted of a mixture of managed and semi-natural stands. The last
cluster of the 6th level (N = 1) was characterised by Cladonia fimbriata, which mainly grows
on coarse woody debris and exposed ground (Kruys et al. 1999), but only seldom on top
of the humus layer. The cluster on the right-hand side mainly contained young, mature
and overmature semi-natural stands.

3.3.3 Summary of the DCA and TWINSPAN analysis

The main results of the DCA ordination is that (i) the semi-natural saplings were located
on the left, (ii) burnt areas in the right corner and (iii) older mesic sample plots in the
middle. The location of older mesic stands is the correlated with the volume of deciduous
trees and dominant height of the trees (Fig. 4). The TWINSPAN classification separated
the same main groups. Cladonia species and Calluna also indicated semi-natural sub-
xeric saplings with a high abundance (Fig. 4 and IV), as in TWINSPAN. Many liverwort
species growing on CWD (III) and Bryum capillare were located near to each other in the
right corner in the DCA ordination. Brachythecium species and also many liverworts were
located in the direction of mesic old stands.
26 Uotila

Fig. 5. TWINSPAN classification of the vegetation data. The indicator species of each
division are given. Numbers after species names give the abundance value of
pseudospecies formation. Numbers in parentheses give the frequencies of the species in
the left- and right-hand clusters, respectively. Mesic sites have been marked with green
and sub-xeric sites with brown symbols. The stage of succession and management class
have been marked with different symbols below.
Results and Discussion 27
28 Uotila

3.4. Patterns of understorey vegetation

3.4.1 Species groups in general

Regardless of the site fertility, the total number of plant species was the greatest in the
semi-natural and managed sapling stands. The greatest difference in the species richness
between managed and semi-natural stands is in the early succession in sapling and young
stands (the mean age about 20 to 50 year). The mean number of dwarf shrubs and lichen
species was twice as high in semi-natural sub-xeric sapling stands as
in managed sapling stands. In the mesic semi-natural sapling and young stands, the
number and coverage of dwarf shrub species were much higher than in managed stands.
The total number of species was relatively similar in semi-natural and managed stands;
the highest in the sapling stands, and decreasing towards older ones (Tonteri 1994). The
colonisation of species in the beginning of succession was faster after clear-cutting than
after fire. The highest number of species is expected to occur at intermediate time spans
during secondary succession (Connell 1978, Huston 1979) and in the intermediate age
since the disturbance (Lahti and Väisänen 1987, Tonteri et al.1990, Nieppola 1992, Tonteri
1994, Collins et al. 1995). The results of this study agree with the intermediate disturbance
hypothesis (III, IV).
Liverworts were more diverse compared to any other group of species in the semi-
natural mesic stands than in managed mesic forests (III). The uneven-aged stand structure
of the sapling and young stage (I, II) provides special conditions with shading and moist
coarse woody debris for liverwort species like Calypgeia muelleriana ( Kvavchenko et al.
2000), Lophozia longiflora (Andersson and Hytteborn 1991, Kruys et al. 1999), Cephalozia
affinis (Söderström 1988, Rassi et al. 2001), Calypogeia neesiana (Kruys et al. 1999),
Calypogeia integristipula and Chiloscyphus profundus (Kruys et al. 1999). Many liverwort
species also live on exposed mineral soil, like Cephaloziella divaricata and Cephalozia
pleniceps. Blepharostoma trichophyllum (Andersson and Hytteborn 1991, Kruys et al.
1999) and Lophozia sudetica. Lophozia longiflora occurred mainly in semi-natural stands,
as was the case for Ptilidium pulcherrinum (III; IV (App.), Kruys et al. 1999). They require
decaying wood and a humid climate for successful growth (Söderström 1988, Andersson
and Hytteborn 1991, Jonsson 1993b).
The occurrence of liverworts, mainly the genera Barbilophozia, Calypogeia, Cephalozia
and Lophozia, seemed to be more diverse in the semi-natural plots on the Russian side
than on the Finnish side (III, IV, App.). In the semi-natural forests, the number of liverwort
species was more than double that in managed stands, except for the young stands.
Cuttings reduce the number of liverworts, especially in sapling and young stands
(Söderström 1988, Kruys et al. 1999). Clear-cuttings also reduce the diversity of liverworts
and increase the coverage of grasses. The shelterwood system and fallen retention trees
 Results and Discussion 29

seem to offer more natural moist and stable conditions for the continuity of liverwort
species (Hannerz and Hånell 1997).
Bryophyte succession on decaying logs and stumps is initiated by liverworts, followed
by more competitive acrocarpous mosses and finally by pleurocarpous mosses
(Krusenstjerna 1945). Bryophytes live in habitats that are more or less dynamic (Söderström
1995), with the availability of resources (light, moisture, nutrients) and the life history strategy
controlling the success of bryophyte species (competition, mortality and abundance)
(During 1992). Epixylic bryophytes especially require a continuity of coarse woody debris
(Andersson and Hytteborn 1991). On the other hand, recolonisation by pleurocarpous
mosses occurred 5-15 years earlier if the canopy had survived a fire (Schimmel 1993).
This study shows that, of the bryophytes, Calypogeia muelleriana, C. sphagnicola,
Lophozia longiflora, Plagiothecium curvifolium and Hylocomium umbratum mainly occur
in natural stands on mesic sites (Kvavchenko et al. 2000, Nitare 2000), with an indication
of a high degree of naturalness (III, App.). Of the vascular plants, Goodyera repens may
represent a tool for indicating the degree of naturalness of the mesic ground vegetation or
of a stand (III, App.). A large number of cup lichens may represent a high degree of
naturalness in the early successional stages of sub-xeric stands (IV, see also Ahti 1977,
Foster 1985). This phenomenon has also been observed nine years after the fire in the
Kitsi study area (I, II, III, and IV) (A. Uotila unpublished material).

3.4.2 Abundance of bryophytes

In the semi-natural stands on mesic sites, the coverage of mosses peaked in the sapling
stands, and decreased towards mature stages (III; IV). On the sub-xeric sites, the highest
values of the moss cover were found in the mature stands. In the managed forests, the
coverage of mosses increased throughout the succession (III; IV). The pioneer species,
Ceratodon purpureus, was almost totally absent from clear-cuttings, and the coverage of
Pohlia nutans was only one third of that in more mature stands. The coverage of Polytrichum
juniperinum was double in the sapling and threefold in the young managed stands
compared to the unmanaged one (see also Mäkipää 2000b, Reinikainen et al. 2000:
Appendix 4). Ceratodon purpureus (18 % of coverage, mesic), Pohlia nutans (10 %, mesic)
and Polytrichum juniperinum (5 %, mesic) invaded the sites immediately after fire (Kujala
1926b, Sarvas 1937, Schimmel and Granström 1996, Bates 2000, Mäkipää 2000b), but
they were gradually replaced by mosses like Hylocomium splendens (III) and Pleurozium
schreberi (III, IV).
For naturally uprooted patches, Jonsson and Esseen (1990) demonstrated the ability
of species such as Pohlia nutans to colonise at an early stage in the succession, but they
may still occur in patches for more than 100 years. Hylocomiun and Pleurozium were
30 Uotila

effective in invading young semi-natural stands in the mesic sites compared to the managed
stands (Mäkipää 2000b). In the mature and overmature/old-growth stands, H. splendens
and P. schreberi showed a similar pattern (III, IV). However, P. schreberi may be more
successful after logging (IV, Nguyen-Xuan et al. 2000, cf. Hannerz and Hånell 1997) than
H. splendens (Tarkhova and Ipatov 1975).
Burning decreases the acidity of the soil and a considerable amount of nutrients are
released after a fire. Bryophytes utilize effectively the available nutrients. On burnt areas
the absorption is probably more effective than on clear-cutting areas, where nutrients are
supplied by precipitation, dust and litter (Oechel and Van Cleve 1986, Bates 2000). More
than 50 years are needed for the nutrient status to return to the level prior to the fire (Viro
1969, Dyrness and Norum 1983, Pietikäinen 1999). For example, Potter et al. (1995) found
that the total cover of Hylocomium splendens was reduced by 50 % within three years on
fertilised plots (N, P, K) compared to unfertilised plots (Viro 1969, Dirkse and Martakis
1992). In general, the severity of disturbances affects the survival and growth of H.
splendens and P. schreberi and other bryophytes (e.g. Jonsson 1993b, Schimmel and
Granström 1996).
The finding that, the of the common mosses, both P. schreberi and H. splendens are
late-successional species (Foster 1985, Reinikainen et al. 2000), does not gain full support
in this study. On the sub-xeric sites, the coverage of P. schreberi increased along with the
maturity of the tree stand regardless of the management (IV). On the mesic sites, the
tendency was the same, but the coverage varied substantially in the mature stands (III).
The abundance of this species is clearly reduced by logging (Nguyen-Xuan et al. 2000,
Jalonen and Vanha-Majamaa 2001, Rees and Juday 2002). Once the forest canopy has
been fully established (30-40 years), H. splendens and P. schreberi become dominant
(Schimmel 1993, O’Neill 2000). Following more effective fire suppression since the 1950s,
the abundance of fire mosses and colonist species has declined with a consequent increase
in pleurocarpous mosses (Reinikainen et al. 2000: Appendix 4).
The uneven-aged structure probably provides conditions that are stable enough for
the formation of long-lasting carpets in semi-natural stands, regardless of small
disturbances in the ground vegetation and tree layers (Bates 2000). On the other hand, H.
splendens has been found to emerge annually from either mature segments or dormant
buds (Økland and Økland 1996), and not from the diaspore or propagule bank. In contrast,
P. schreberi has been reported to emerge from the diaspore or propagule bank after a
floor disturbance (Johnsson 1993a, Økland 1995). The explanation for the higher mortality
of H. splendens compared to P. schreberi in early succession could be that the latter
species has many survival strategies.
Dicranum majus (III) was most common in the mature semi-natural stands. Mäkipää
(2000b) found that a stand age of 100 years was associated with the highest occurrence
 Results and Discussion 31

of D. majus. This is well in line with the result obtained in this study for semi-natural
stands, but not for managed stands. Regardless of the phase of succession, Dicranum
polysetum and Dicranum scoparium were successful in the managed stands compared
to the semi-natural ones. This result is in agreement with the findings of Söderström
(1988) concerning the occurrence of D. scoparium in old managed stands (at an age of 50
years for spruce and 280 years for pine) and old-growth forests (at an age of 300-350
years). The abundance of D. polysetum and D. scoparium has increased since the 1950s
due to management, as Mäkipää (2000b) reported on the basis of the results of the National
Forest Inventories. In contrast, the coverage of Dicranum fuscences, which mainly lives
on decaying wood, has decreased (Reinikainen et al. 2000: Appendix 4).
Many Brachythecium species like B. rutabulum and B. starkei prefer sites characterised
by an abundance of litter (Söderström 1988, Bates 2000). In this study, seven
Brachythecium species were identified, and all of them were absent from clear-cuttings.
However, these species were common on the sites representing young managed saplings.
This finding is in line with the results from the National Forest Inventory, which show an
increasing occurrence of Brachythecium since the 1950s (Mäkipää 2000b). For example,
the abundance of Brachythecium starkei was greater in managed than in semi-natural
stands, excluding managed overmature stands, as reported by Söderström (1988).
Interestingly, some Brachythecium species seem to be rather successful after nitrogen
fertilisation (Mäkipää 2000b). On the other hand, Brachythecium species may survive the
burning in the patches typical of burnt areas.

3.4.3 Abundance of lichens

Cladina rangiferina and C. arbuscula were the most abundant in the sapling stage after
canopy fire (IV). The coverage of these species decreased sharply in the young stands,
but then increased clearly along with the increasing maturity of the semi-natural stands.
The occurrence of lichens is clearly related to the succession gradient on dryish heaths
(IV). The density of living trees affected the abundance of Cladina mitis, Cl. rangiferina and
Cl. stellaris in the semi-natural sub-xeric sites, but not in managed ones (I, IV). Cl. rangiferina
and Cl. stellaris had higher abundances in the areas burnt only lightly, and Cl. mitis was
abundant in open areas that had experienced severe fires (Arseneault 2001).
Lichens are prominent in the secondary succession of boreal forests after natural
disturbances (Coxson and Marsh 2001). Ahti (1977) divided lichens into the following
chronosequence: 1. Bare soil (1-3 yrs after fire); 2. Crustose lichen stage, (3-10 yrs); 3.
Cup lichen stage (10-50 yrs); 4. First reindeer lichen stage (30-120 yrs); 5. Second reindeer
lichen stage (80-120 yrs). Based on Ahti (1977), it could be expected that lichens would
32 Uotila

have a higher species number in the semi-natural sapling stands compared to the managed
sapling stands, as was found in this study (IV) and reported previously in several papers
(Foster 1985, Longton 1992 and Nguenyen-Xuan 2000). On the other hand, the number
and the coverage of lichens were lower in managed dryish heaths (IV), and ground lichens
were virtually absent from mesic spruce-dominated stands (IV) (Esseen et al. 1997,
Humphrey et al. 2002).
Reindeer lichens suffer from cuttings (Bråkenhielm and Liu 1998), and the occurrence
of Cladonia species may be connected with the availability of different nutrients. For
example, Cladonia species seem to disappear after nitrogen fertilisation (van Dobben
1993, Vagts and Kinder 1999), probably because ammonium ions (NH4+) are toxic to these
species (Nash III 1996). Lichens are the most successful in extreme microclimatic conditions
where there are virtually no other plant species competing for resources (Kershaw 1977,
Kershaw 1985, Sulyma and Coxson 2001).

3.4.4 Abundance of dwarf shrubs

The coverage of dwarf shrubs in sub-xeric stands increased along with the maturity of the
tree stands regardless of the management. This was not the case in the mesic sites,
where the coverage of dwarf shrubs culminated in the semi-natural stands representing
the sapling stage (III, IV).
Calluna invades with the help of seeds dispersed outside the site or from seeds buried
in the soil (seed bank). Calluna is the most abundant in semi-natural sapling stands (Kujala
1926a, Sarvas 1937, Granström and Schimmel 1993, Valbuena and Trabaud 2001). Another
reason for invasion in the sapling stage could be changes in the amount of available
calcium in the soil (Uggla 1957, Viro 1969, Dyrness and Norum 1983, Mälkönen and Levula
1996), which increases considerably after fire compared to the situation after clear-cutting.
Leake and Read (1989) reported that the growth of mycorrhizal-infected Calluna was
significantly higher than noninfected Calluna at sites with lower calcium concentrations.
Regardless of the site type, the coverage of Vaccinium vitis-idaea was larger in the
sapling and young stands representing managed forests than in the semi-natural ones of
the development phases (IV). In the mature stands, the coverage was fairly same regardless
of the management. Regarding V. myrtillus, the coverage culminated in the semi-natural
stands representing the sapling and young stages in the mesic sites. In the sub-xeric
stands the coverage of V. myrtillus increased along with the increase of the maturity of the
tree stands regardless of the management. However, management reduced the coverage
of V. myrtillus in the mature stands representing the sub-xeric sites (IV) as found in some
previous studies (Atlegrim and Sjöberg 1996, Salemaa 2000, Reinikainen and Salemaa
 Results and Discussion 33

2000, Reinikainen et al.2000) with respect to the logging intensity. This was also the case
for the managed young stands on the mesic sites (III) (Atlegrim and Sjöberg 1996, Salemaa
2000, Reinikainen and Salemaa 2000, Reinikainen et al.2000, Bergstedt and Milberg 2001,
Deal 2001. Similarly, Empetrum nigrum seems to suffer from fire, and also to recover
slowly after cuttings (IIII; Kujala 1926a, Sarvas 1937, Foster 1985).”

3.4.5 Abundance of grasses

In the managed stands the abundance of grasses was the highest in the early succession,
i.e. on clear-cuttings and in the sapling stands (III, IV). Grasses are able to compete with
other vegetation after cuttings on moderately “undisturbed” ground, but not so well on
burnt land (Kershaw 1985). Management seemed to favour Deschampsia flexuosa the
most (III, IV). The rhizomes of Deschampsia flexuosa survive in cuttings, and the high
availability of nutrient and light on such sites are the reasons for the overwhelming success
of this species (Foggo 1989, Högbom et al 2002). The relative importance of disturbance
for the success of this species is different after clear-cutting with soil preparation than
after a ground or canopy fire. Clearly, grasses occupy rapidly clear-cutting areas (III, IV),
whereas the abundance of grasses and herbs in semi-natural stands is low (Kujala 1926a,
1926b, Sarvas 1937, Ferm and Pohtila 1977). The coverage of Deschampsia flexuosa has
increased since the 1950s due to the change in forest age structure (Mäkipää 2000a).

3.5. Evaluation of the results

This study utilises the temporary sample plots representing the different phases of forest
succession. This approach is widely applied, and the only way to build the successional
series for forest development. Unfortunately, this will result in variability, which is not
related to the successional process. An attempt was made to avoid these problems through
careful selection of the study sites so that the site type is the same through successional
series in both the unmanaged and managed stands. When doing this, the forest site
types of Cajander (1949) were applied in order to make a distinction between the sites of
varying fertility. For the problems involved in constructing a successional series based on
temporary plots, see Sirén (1955) and Halpern and Spies (1995).
The managed stands located in Finland were compared to the semi-natural stands
located either in Finland or Russia. Both areas are characterised by similar weather and
land use history in terms of slash-and-burn cultivation, although it has been more intensive
in Finland than in Russia. About half of the data of the young semi-natural stands (stages
34 Uotila

II and III) are from Finland and half from Russia. Similarly, the mature and old-growth
stages partly include sample plots from the Russian side, except for the mesic old-growth
forests that were located only in Finland. This mixture of sample plots representing different
locations decreases the effect of location, and eliminates systematic error in sampling (I,
II). On the other hand, in young successional stages the understorey vegetation is more
random than in closed, older successional stages (Cajander 1949, Nieppola 1986).
However, the vegetation of semi-natural sub-xeric saplings in Russia deviated clearly
from that in Finland (Fig. 5). This difference most probably reflects the differences in land
use history. The semi-natural vegetation deviated in the same way from the managed one
regardless of the location (Fig. 4).
The results may be also biased by differences in forestry practices (cuttings and
management). The differences induced by cuttings and management may be highlighted
with the help of the management points representing the sum of the treatments subjected
to the stands as presented in the article II (III, IV Table 1) as regards the evaluation of
management history of the study plots. It appeared that a part of the semi-natural stands
had been subjected to light selection fellings, and the managed stands were clearly more
intensively cut and managed. Therefore it is concluded that the results reliably represent
the vegetation patterns in the study area, which is modified by natural and human-induced
disturbances. In some previous studies, the impacts of land use and management history
are omitted, and emphasis placed on the structural and species differences between
mature and old-growth successional stages representing managed and natural forests
(e.g. Siitonen 2001, Table 1). For information about the impacts of land use and management
history on forest succession see, however, Östlund et al. (1997), Linder (1998a, b) and
Pitkänen et al. (2003).
The history of land use and management of the forests in the research areas is variable,
but most probably clear cuttings have not initiated the succession behind the forests now
in the mature phase. The initial stands are likely representatives of those after light selective
cutting, which has continued throughout the succession. Consequently, it was not expected
that forest management of 20th century with clear cuttings will produce forests developing
in every detail to their full maturity following the patterns observed in this study. Especially,
on the mesic sites grass vegetation is enhanced substantially after clear cutting. This will
probably increase the grass vegetation throughout the succession. This kind of pattern
was found in this study, but it cannot directly generalise to represent the pattern after the
clear cutting. The only complete and fully satisfying solution for this methodological problem
would be a series of permanent study plots that are monitored over a very long time
period (see Sernander 1936). This is quite naturally impossible in practice.
The structural differences between the semi-natural and managed stands in the same
successional stage were also analysed with help of the Turkey’s test. This test showed
 Results and Discussion 35

the general differences in succession dynamics, but no differences between the managed
and semi-natural successional stages except for the coverage of grasses in mesic sites
and the coverage of bryophytes and grasses in sub-xeric sites. This difference between
semi-natural and managed stands is to be expected, as demonstrated earlier by
Reinikainen et al. (2000) (III, IV). This result is in line with the land use and management
history of the semi-natural and managed stands investigated in this study as appears
from Table 1 (III, IV, see also II). The values of management scores are higher for old
managed forests than for old semi-natural ones.
36 Uotila

4. CONCLUSIONS

Natural plant communities represent a mosaic of structural phases, and naturalness must
be seen as continuous variables with many dimensions. The present results strongly
suggest that natural stand dynamics have not been the only factor modifying the current
structure of the mature forests in Finnish nature reserves and national parks (II). Likewise,
old, selectively logged (managed) forests have some characteristics of near natural stands
and features of naturalness. Cuttings and management have decreased the abundance
of liverwort species in mesic sites and lichens in sub-xeric sites, and the coverage of
dwarf shrubs such as Calluna vulgaris in sub-xeric stands and Vaccinium myrtillus in
mesic sites (III, IV). However, there was large variability in the coverage of different species,
which hampers clear conclusions. Cuttings and management have increased the coverage
of grasses in both mesic and sub-xeric sites, and the coverage of mosses in sub-xeric
sites (III, IV).
Research and conservation efforts have mainly been focused on old-growth forests in
order to preserve these habitats and their threatened species. This choice is partly
influenced by the presence of coarse woody debris, which maintains the diversity of
habitats and the consequent diversity of species. However, the young successional stages
of natural or semi-natural forests are also valuable for conservation (I; II, III, IV, Similä
2002, Similä et al. 2002, 2003), because they represent an uneven-aged stand structure
and the presence of coarse woody debris. They probably maintain conditions that are
suitable for lichens and liverwort species or for the regeneration of dwarf shrubs (III, IV)
and provide variable habitats after fire. This implies that natural forests should also include
different successional stages (I, III, IV). Therefore, the focus in the restoration of conservation
areas should be on the dynamics of forest landscape in relation to the habitat requirements
of threatened and rare species, as well as general species at the stand level (III, IV).
The future research needs require permanent study plots and long-term monitoring of
the re-vegetation strategies of representative species in young natural stands, e.g. of the
main dwarf shrubs, grasses, mosses and lichens, compared to the strategies in managed
stands in relation to fertility. Furthermore, there is a need to enhance our understanding of
regeneration dynamics after forest fires, storms and clear-cuttings. Obviously, the history
of land use and management is important for understanding “natural” biodiversity when
attempting to optimise forest management to maintain biodiversity. No uniform silvicultural
instructions, designed to maintain the diversity of structures and species in forests, can
be made for the whole of Finland owing to variations in the management history in different
parts of the country.
The common practice in Finland of leaving groups of trees in regeneration cuttings
creates a residual tree layer in the areas. This may later increase the volume of dead
wood, which is urgently needed throughout the succession in order to provide habitats
 Conclusions 37

for the species closely associated with the occurrence of dead wood debris (I). This, in
addition to natural regeneration, would also further increase the number of tree storeys in
the older successional stages. Fires and natural disturbances play a key role in achieving
a closer resemblance to natural succession dynamics in managed forests, because they
are of primary importance for forming coarse woody debris, uneven-aged stand structure
(I, II) and microhabitat patches for the understorey vegetation (III, IV).
38 Uotila

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