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ROUTLEDGE HANDBOOK OF
FOREST ECOLOGY
This comprehensive handbook provides a unique resource covering all aspects of forest ecology
from a global perspective. It covers both natural and managed forests, from boreal, temperate,
subtropical and tropical regions of the world. The book is divided into seven parts addressing
the following themes:
• forest types
• forest dynamics
• forest flora and fauna
• energy and nutrients
• forest conservation and management
• forests and climate change
• human impacts on forest ecology.
While each chapter can stand alone as a suitable resource for a lecture or seminar, the complete
book provides an essential reference text for a wide range of students of ecology, environmental
science, forestry, geography and natural resource management. Contributors include leading
authorities from all parts of the world.
Kelvin S.-H. Peh is Lecturer in the Faculty of Natural and Environmental Sciences, University
of Southampton, and also Visiting Fellow in the Department of Zoology, University of
Cambridge, UK.
Richard T. Corlett is Professor and Director of the Centre for Integrative Conservation,
Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Yunnan, China. He
was previously a professor at the National University of Singapore and the University of Hong
Kong.
Typeset in Bembo
by Saxon Graphics Ltd, Derby
CONTENTS
Contributors xi
1 Introduction 1
Kelvin S.-H. Peh, Yves Bergeron and Richard T. Corlett
PART I
The forest 5
2 Boreal forests 7
Jean-Pierre Saucier, Ken Baldwin, Pavel Krestov and Torre Jorgenson
4 Subtropical forests 46
Richard T. Corlett and Alice C. Hughes
5 Tropical forests 56
Lindsay F. Banin, Oliver L. Phillips and Simon L. Lewis
6 Managed forests 75
Jürgen Bauhus and Patrick Pyttel
v
Contents
PART II
Forest dynamics 91
PART III
Forest flora and fauna 183
vi
Contents
PART IV
Energy and nutrients 307
PART V
Forest conservation and management 369
vii
Contents
PART VI
Forest and climate change 471
33 Fire and climate: using the past to predict the future 473
Justin Waito, Martin P. Girardin, Jacques C. Tardif, Christelle Hély,
Olivier Blarquez and Adam A. Ali
PART VII
Human ecology 557
viii
Contents
Index 634
ix
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CONTRIBUTORS
Ken Baldwin; Forest Ecologist, Natural Resources Canada, Canadian Forest Service, Canada
Andrea Battisti; Professor, Department of Agronomy, Food, Natural Resources, Animals and
the Environment, University of Padova, Italy
xi
Contributors
Shabtai Cohen; Senior Research Scientist, Institute of Soil, Water and Environmental
Sciences, Agricultural Research Organization (ARO), Volcani Center, Israel
Lluís Coll; Research Scientist, Forest Sciences Centre of Catalonia (CTFC), Spain
Barry Cooke; Research Scientist, Northern Forestry Centre, Natural Resources Canada,
Canadian Forest Service, Canada
Lee E. Frelich; Director, Center for Forest Ecology, University of Minnesota, USA
xii
Contributors
Christelle Hély; Director of Studies, École Pratique des Hautes Études, Institut des Sciences
de l’Évolution - Montpellier, UMR 5554, France
Dave Kendal; Ecologist, Australian Research Centre for Urban Ecology, Royal Botanic
Gardens Victoria and University of Melbourne, Australia
David Lamb; Honorary Research Fellow, School of Agriculture and Food Sciences and
Centre for Mined Land Rehabilitation, University of Queensland, Australia
Mark J. McDonnell; Director, Australian Research Centre for Urban Ecology, Royal Botanic
Gardens Victoria and Associate Professor, University of Melbourne, Australia
Yadvinder Malhi; Professor, Environmental Change Institute, School of Geography and the
Environment, University of Oxford, UK
Daniel Markewitz; Professor, Warnell School of Forestry and Natural Resources, University
of Georgia, USA
xiii
Contributors
David Paré; Research Scientist, Laurentian Forestry Centre, Natural Resources Canada,
Canadian Forest Service, Canada
John A. Parrotta; Program Leader for International Science Issues, U.S. Forest Service
Research and Development, USA
Kelvin S.-H. Peh; Lecturer, Centre for Biological Sciences, University of Southampton and
Visiting Fellow, Department of Zoology, University of Cambridge, UK
Emma J. Pharo; Senior Lecturer, School of Land and Food, University of Tasmania, Australia
xiv
Contributors
Hazel K. Smith; Research Fellow, Centre for Biological Sciences, University of Southampton,
UK
Jeffrey A. Stratford; Associate Professor, Department of Biology and Health Sciences, Wilkes
University, USA
Leho Tedersoo; Senior Researcher, Natural History Museum and Botanical Garden, Tartu
University, Estonia
Edgar C. (Ed) Turner; Academic Director and Teaching Officer in Biological Sciences,
Institute of Continuing Education and Post-doctoral Researcher, Department of Zoology,
University of Cambridge, UK
Stephen M. Turton; Professor, Centre for Tropical Environmental and Sustainability Science,
James Cook University, Australia
xv
Contributors
Justin Waito; Researcher, Centre for Forest Interdisciplinary Research (C-FIR), Department
of Biology, The University of Winnipeg, Canada
Simon Willcock; Research Fellow, Centre for Biological Sciences, University of Southampton,
UK
xvi
1
INTRODUCTION
Kelvin S.-H. Peh, Yves Bergeron and Richard T. Corlett
Forests are stupendous systems. Since the first trees appeared on Earth in the Late Devonian,
390 million years ago, the complex three-dimensional structure of forests has supported the
majority of terrestrial species on Earth and this is still true today. We ourselves evolved from a
predominantly forest lineage, but the first humans occupied more open habitats and this non-
forest origin is reflected in our ambiguous relationship with forests. On the one hand, this
relationship until now has been largely destructive, with forests valued most as a source of land
for cultivation, and for timber and bushmeat. On the other hand, on a crowded planet we
increasingly value the services that forests can provide: clean water, erosion control, and the
amelioration of local, regional and global climates. Forests are also important for recreation, and
as sources of artistic and spiritual inspiration. Reconciling these incompatible objectives will
require both a better understanding of forest ecology and a wider awareness of the multiple
values of forests. We hope this book will contribute to both these aims.
Our forest systems today – unfortunately – are facing a suite of global challenges. Deforestation
and forest degradation, biological invasions, excessive and often illegal harvesting of forest
products, atmospheric pollution and climate change – to name only the biggest challenges – are
impacting our forests in a profound way, and we are still learning how these systems are actually
coping. Deforestation has emerged as one of the most damaging problems, as large areas are
cleared and converted into agricultural land and livestock ranches to feed an ever-growing
population, particularly in the tropics. Large-scale production of biofuels poses a potential
additional threat. Climate change, which results in increasingly extreme weather, has also
emerged as equally detrimental, leaving its mark on our forests in diverse ways – from erosive
floods and deadly droughts to vanishing coastal mangrove habitats. Worse still, these
environmental challenges seldom act alone, but also occur together, simultaneously and interact
to further aggravate the problems, thereby putting an unprecedented pressure on the forest
biodiversity and function. Policy-makers involved in ongoing international agreements, such as
the Convention on Biological Diversity, and practitioners in international programmes, such as
the United Nations’ collaborative initiative in reducing emissions from deforestation and forest
degradation (REDD), are seeking practicable solutions. Again, there is a need to provide to a
wider audience reliable information on forests, their dynamics, biodiversity and responses to
human disturbance and climate change, as well as the applications of ecology in the management
and mitigation of these global challenges.
1
Kelvin S.-H. Peh, Yves Bergeron et al.
This handbook aims to act as a state-of-the-art summary of our current knowledge of forest
ecology. It draws on the expertise of a varied international team of authors, many of whom are
experts in their respective field, or practitioners with rich experience in forestry. It aims to be
an informative, up-to-date resource on the literature on forests and their ecology under
environmental change. The handbook does not promote any particular viewpoint, management
practice or conservation approach. Instead, it covers a broad range of subjects subsumed under
the realm of forest ecology, and offers a comprehensive overview for each of these topics. Most
chapters aim for a global coverage as much as practicably possible, but others focus on the
region where the subject is most relevant or has received most attention. By presenting each
topic across different geographical areas – or at least those biomes whose processes are
significantly different – and cross-comparing them as appropriate, we aim to give readers a
unique perspective.
The work is grouped around seven parts. The first section of the handbook, ‘The forest’, is
intended to help define different major forest biomes loosely according to latitudinal belts. The
forest types covered are: boreal forests, northern temperate forests, subtropical forests, tropical
forests and managed forests. These chapters provide an introductory overview setting out
definitions, scopes and different forest types within each major forest biome. Thus, Part I
provides a primer on the forest systems from five different forest biomes which provide the
framework for understanding the other themes. The list of forest biomes covered is not
exhaustive, in that we do not cover some important topics such as the southern temperate
forests and the mountain forests. Nevertheless, the vast majority of the forest biomes are
discussed in this section.
Part II, ‘Forest dynamics’, includes a series of chapters that explore the impacts of different
disturbances – namely insects, fire and strong winds – on forests. The common theme in this
section is that the forest systems often show resistance and resilience to perturbation and
apparently are able to recover from disturbances. However, disturbances may exceed a critical
level whereby systems rapidly and irreversibly collapse, and it is important to understand the
basis for such tipping points, the circumstances under which such phenomena may be reached
and how to prevent reaching them. This section also includes chapters that discuss how the
biological factors, scaled from genes up to a community of hundreds of interacting species,
shape and change the forests.
Part III is intended to showcase the rich biodiversity of the forests. Natural history is needed
to support the understanding of major ecological processes. This section therefore concerns the
ecology of important taxa such as lianas, vascular epiphytes, bryophytes, lichens, insects,
mammals and birds. It also includes a brief introduction to the microbial pathogens and insect
pests – the major agents of biological disturbances in both natural and managed forests. As
forests harbour a huge variety of organisms, we acknowledge that the taxa discussed in this
section cannot be comprehensive; for example, reptiles and amphibians are not covered.
Nevertheless, this section ends with a concluding chapter that provides a global perspective on
forest biodiversity, evaluating our current knowledge of species diversity in our global forests.
In Part IV, the chapters focus on the complexity of some important ecological functions that
ensure energy and nutrients are acquired, utilised and recovered in a clockwork fashion. Any
missing components – for instance, mycorrhizal associations – will dramatically jeopardise the
net primary productivity. Likewise, any processes that are slowed down will be further
diminished by a negative feedback mechanism. For example, a slower rate of nutrient cycling
will substantially reduce the net primary productivity, in turn lowering plant nutrient availability
and exacerbating the effects of nutrient limitations. These chapters invite the reader to appreciate
the fragility of our forest systems. This section therefore builds an ecological perspective of
2
Introduction
sheer complexity in the engineering of the forests as whole systems that provide an essential
foundation supporting all life.
The chapters in Part V examine the ecology behind some of the cross-cutting anthropogenic
threats and some conservation approaches that inform policy and management practices. All
subjects covered in these chapters either play an important ecological role in restoring the forest
systems or are significant challenges that place obstacles in the path of preserving their
biodiversity and function. We admittedly consider these issues in separate chapters, rather than
simultaneously, although our ability to manage the forests will increasingly require an
understanding of the synergistic effects of multiple challenges.
Climate change is the focus of Part VI. This part includes a series of chapters that explore
the impact of different climate change phenomena, such as fire and drought. But the list of
major global threats due to climate change is not exhaustive, for we do not cover topics such
as the rising sea level and its impact on coastal and insular forests, which can be profound, or
the impact of increased heavy precipitation events over forested areas. As in the previous
section, we consider these threats caused by climate change one at a time, but the last chapter
of this section shows how large-scale modelling approaches can help provide an integrated
understanding of multiple global threats.
The final part of the handbook examines the direct use of forests by people and its impact
in a wider context of maintaining or sustaining the forests’ capacity to provide ecosystem
services. The chapters within this section discuss some major services which the forests
contribute – such as the provision of timber and non-timber forest products, and areas for
productive cultivation and nature-based recreation, which includes urban tree-dominated
‘green spaces’ and forests for recreational hunting. Together these chapters summarise the
connections between human well-being and the health of our global forests.
Like all works of this kind, the handbook represents a balance between the need to
disseminate further knowledge, and the imperative of not missing the narrative in a welter of
details. The intent for this volume is to be an authoritative text, yet at the same time appealing
to the layperson interested in an introduction to forest ecology, as well as providing graduate
students with a comprehensive collection of current research for further examination and
discourse on the subject. Our aim is also to narrate the ecology behind some of the current
approaches and trends in forest management to conservation practitioners. We are thankful to
every contributor to this volume for sharing their cutting-edge knowledge. Lastly, we are also
grateful to a team of outstanding anonymous reviewers for their time and hard work. For their
concerns, criticisms, feedback and suggestions have also helped us bring this handbook to a
higher level.
3
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PART I
The forest
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2
BOREAL FORESTS
Jean-Pierre Saucier, Ken Baldwin, Pavel Krestov and Torre Jorgenson
The boreal biome is one of the largest forested biomes on Earth and forms a circumpolar belt
of forests and woodlands between the treeless arctic zone and the temperate zone. It is typically
characterized by a cold continental climate with relatively short, mild summers and long, cold
winters. About one-third of the biome occurs in the zone of permafrost (Brown et al. 1997).
Boreal forests and woodlands represent approximately 33 per cent of the world’s forested area
(FAO 2001). Boreal forests are dominated by a relatively few, primarily coniferous, genera
(Picea, Abies, Larix, Pinus) that are adapted to cold temperatures, low nutrient conditions and
recurrent stand-replacing disturbance. Short growing seasons and cold, acidic mineral soils
under conifer canopies result in extensive feathermoss carpets on upland sites. Sphagnum
mosses occupy landscape positions with permanently high water tables, resulting in acidic
organic soils that often develop into peatlands. On both upland and lowland sites, decomposition
rates are slow and nutrient cycling is typically restricted to the upper soil layers where oxygen
and increased temperatures support microorganism and fungal metabolism. Understory
vegetation of boreal forests is also dominated by a relatively few botanical families, especially
the Ericaceae (the heath or blueberry family) which is adapted to cold, nutrient impoverished
habitat conditions. With proximity to the oceans, winters become milder and summers cooler;
snow covers the ground for longer periods and the growing season is shortened. In these
oceanic boreal climates, conifers are generally absent and are replaced by birch (Betula spp.),
alder (Alnus spp.) or ericaceous shrublands that can tolerate such harsh conditions.
7
Jean-Pierre Saucier, Ken Baldwin et al.
Figure 2.1 Extent of boreal forests and woodlands around the globe, with floristic subdivisions and
non-forest boreal vegetation zones
to the Pacific coast. Boreal vegetation also covers some islands in the Atlantic (Iceland, Faroe
Islands and the southernmost part of Greenland) and in the Pacific (Kuril and Aleutian island
chains). Closed forest covers 76 per cent of the boreal zone, while commercially exploitable
closed forest occupies about 53 per cent of the total area (Table 2.1).
The northern limit between the boreal zone and the arctic zone is the continental treeline
(i.e., the edge of the habitat in which trees are capable of growing). Moving northward from
closed forests through open forests and woodlands, climatic and/or site conditions eventually
become too harsh (cold, dry, windy, infertile) to support tree growth. Trees are replaced by
low or prostrate shrubs, especially dwarf ericaceous, birch and willow species, together with
herbs, mosses and lichens as forest and woodland gradually changes to arctic tundra.
The southern boundary of the boreal zone is less obvious as it is based on vegetation
physiognomy, species composition and ecosystem dominance on the landscape. This boundary
represents a gradual shift of forest composition where, moving southward, thermophilous
species replace the boreal species (Brandt 2009). It is usually marked by a change from a
coniferous dominated landscape, associated with a cold climate, to a mixed forest landscape
with a milder climate (see next section). In certain areas, with soils of variable fertility and very
dry climate, the boreal forest is replaced by steppe at its southern margin.
8
Boreal forests
Table 2.1 A
rea of boreal forests and woodlands by country and proportion of the country in closed
forest or exploitable closed forest (adapted from Kuusela 1992)
Area of forest and other wooded land within the boreal zone (million km2)
Transition zones
Between the boreal zone and the adjacent vegetation zones, many authors recognize transition
zones.
Between the boreal zone and the arctic zone, the transition is called hemiarctic, or subarctic.
This ecotone is characterized by a landscape matrix of woodlands and treeless barrens where
rare stands of trees grow in sheltered locations. Accumulated snow usually protects the trees
during the coldest months. These tree stands are often embedded in krummholz vegetation.
Authors agree that the hemiarctic is part of the boreal zone, since its flora retains species that
are at the northern limit of their ranges, such as the tree species.
Between the boreal zone and the temperate zone, the transition is called hemiboreal. In the
hemiboreal, boreal species form mixed forest types with species that are less cold tolerant and
are at the northern limit of their ranges. For example, in eastern North America, balsam fir
(Abies balsamea1) and white spruce (Picea glauca), characteristic of the closed boreal forest, form
mixed stands with yellow birch (Betula alleghaniensis) and sugar maple (Acer saccharum). In eastern
Asia, a mix of broadleaved tree species (Acer mono, Betula costata, Fraxinus mandshurica, Quercus
mongolica) and conifers (Pinus koraiensis) marks the ecotone between the boreal and temperate
zones. Some authors classify the hemiboreal as a part of the boreal zone. Considering that
temperate species found in this transition zone are at the northern limit of their ranges and
extend far south into the temperate zone, hemiboreal vegetation fits better into the temperate
zone (Saucier 2008). This treatment is logically consistent with classification of the hemiarctic
vegetation into the boreal zone.
1 For vascular plants, we follow the nomenclature of Flora of North America (1993+) for North America and
Cherepanov (2007) for Europe and Asia.
9
Jean-Pierre Saucier, Ken Baldwin et al.
In eastern North America, some high elevation forests, embedded into the northern
temperate zone, share species such as Abies balsamea, Picea glauca and Betula papyrifera with the
boreal zone further north, but also harbour Picea rubens, a more temperate species. This is
the case over 1000 m elevation in New England. On a few of the highest mountain peaks, over
1300 m elevation, numerous arctic species occur as disjunct populations.
In the Rocky Mountains of western North America, a similar situation can be observed.
Boreal species such as Pinus contorta, Picea glauca and Abies lasiocarpa are replaced at higher
elevations southwards by temperate species of pine (e.g., Pinus edulis), fir (e.g., Abies concolor),
spruce (e.g., Picea engelmannii) and juniper (e.g., Juniperus osteosperma).
In Asia, boreal-like forests occur in the mountains of the Japanese archipelago, where they
are dominated by Abies veitchii, Abies mariesii and Abies homolepis. Likewise, forests of Abies
koreana occupy the highest elevations of the mountains in the southern Korean peninsula
(Nakamura and Krestov 2005). Further south, Abies kawakamii occurs above 3000 m in the
mountains of northern Taiwan. All of these forests include some boreal species in the understory,
but most of their flora is composed of temperate species.
Bogs and poor fens occurring in temperate regions contain boreal species that are
characteristic of cold, acidic, nutrient-poor peatlands. In North America, Picea mariana, Larix
laricina and Sphagnum spp. are typical of these oligotrophic wetlands while in Asia, Picea glehnii
and several Larix species are found in these habitats.
10
Boreal forests
Table 2.2 C
limatic values for bioclimates within the boreal macrobioclimate (from Rivas-Martínez et al.
2011)
11
Table 2.3 Dominant boreal tree species by continent and floristic subdivisions of the boreal zone
Picea glauca (White spruce) Picea glauca (White spruce) Picea glauca (White spruce)
Picea mariana (Black spruce) Picea mariana (Black spruce) Picea mariana (Black spruce)
Larix laricina (Tamarack) Larix laricina (Tamarack) Larix laricina (Tamarack)
Betula neoalaskana (Alaska Pinus contorta (Lodgepole pine) Pinus banksiana (Jack pine)
paper birch) Pinus banksiana (Jack pine) Abies balsamea (Balsam fir)
Populus tremuloides (Quaking Abies lasiocarpa (Subalpine fir) Betula papyrifera (Paper birch)
aspen) Abies balsamea (Balsam fir) Betula pubescens (Downy birch)
Populus balsamifera (Balsam Betula papyrifera (Paper birch) Populus tremuloides (Quaking aspen)
poplar) Betula nana (Dwarf birch) Populus balsamifera (Balsam poplar)
Populus tremuloides (Quaking aspen)
Populus balsamifera (Balsam poplar)
North European
Picea obovata (Siberian spruce) Picea obovata (Siberian spruce) Larix gmelinii (Dahurian larch)
Abies sibirica (Siberian fir) Abies sibirica (Siberian fir) Picea obovata (Siberian spruce)
Larix sibirica (Siberian larch) Larix sibirica (Siberian larch) Abies sibirica (Siberian fir)
Pinus sibirica (Siberian pine) Pinus sibirica (Siberian pine) Larix sibirica (Siberian larch)
Pinus sylvestris (Scots pine) Pinus sylvestris (Scots pine) Larix cajanderi (Cajander’s larch)
Betula pendula (Silver birch) Betula pendula (Silver birch) Pinus sibirica (Siberian pine)
Populus tremula (Eurasian Betula platyphylla (Asian white birch) Pinus sylvestris (Scots pine)
aspen) Populus tremula (Eurasian aspen) Betula platyphylla (Asian white birch)
Populus tremula (Eurasian aspen)
Larix gmelinii (Dahurian larch) Larix cajanderi (Cajander’s larch) Picea jezoensis (Jezo spruce)
Pinus sylvestris (Scots pine) Pinus pumila (Siberian dwarf pine) Abies nephrolepis (Manchurian fir)
Pinus sibirica (Siberian pine) Pinus sylvestris (Scots pine) Abies sachalinensis (Sakhalin fir)
Picea obovata (Siberian spruce) Betula platyphylla (Asian white birch) Larix cajanderi (Cajander’s larch)
Abies sibirica (Siberian fir) Populus tremula (Eurasian aspen) Betula ermanii (Erman’s birch)
Larix sibirica (Siberian larch) Betula platyphylla (Asian white birch)
Betula pendula (Silver birch) Pinus pumila (Siberian dwarf pine)
Populus tremula (Eurasian aspen) Populus tremula (Eurasian aspen)
Boreal forests
Alaska–Yukon boreal
The Alaska–Yukon boreal is dominated by mixed forests that range from early to late
successional stages due to the prevalence of fire disturbance. While open to closed Picea glauca
forests represent the potential natural vegetation on mesic slopes, Betula neoalaskana–Populus
tremuloides forests, and spruce–birch–aspen mixed forests are intermixed in a patchy mosaic of
differing aged stands, with the three dominant tree species often present in all successional
stages (Viereck et al. 1992). The Alaska–Yukon boreal extends from western Alaska, where the
forest grades into arctic tundra and boreal shrublands and heaths, to the western edge of the
North American Cordillera in Yukon Territory. Latitudinally, it extends from the southern
Brooks Range to the northern side of the Alaskan coastal mountains. It is differentiated from
the West–Central North American boreal mainly by the presence of Betula neoalaskana and the
absence of Pinus contorta var. latifolia and Abies lasiocarpa. Generally, the understory contains
species that are characteristic of alpine areas or northern boreal latitudes (e.g., Vaccinium
uliginosum, Empetrum nigrum, Betula nana, Arctous rubra and Nephroma arcticum). Forests are
typically found up to 900 m in elevation.
The Alaska-Yukon boreal comprises a diversity of forest types related to topographic
position, soil moisture, nutrients and permafrost, and disturbance by fire and thermokarst
activity (Van Cleve et al. 1983, Chapin et al. 2006). In upland permafrost-free areas, vegetation
succession after fire has several stages, including herb stage, shrub and sapling stage, deciduous
forest (Betula neoalaskana, Populus tremuloides) after 30 to 130 years, and coniferous forest (Picea
glauca) after 100 to 250 years. On permafrost-affected soils on north-facing slopes and lowlands,
coniferous woodlands (Picea mariana, Larix laricina) predominate and are highly susceptible to
fire. Thermokarst is prevalent in permafrost-rich lowlands and creates a variety of non-forested
ecosystems. In the northern portion of the region, coniferous woodlands (Picea glauca, Betula
nana) predominate, while in the southern portion ferns and herbs (Gymnocarpium dryopteris,
Dryopteris dilatata) are abundant in the understory and other shrub species (Salix barclayii, Alnus
viridis ssp. sinuata) predominate.
The non-forested region of the North Pacific, including southwest Alaska, Aleutian Islands,
southeastern Kamchatka, and northern Kuril Islands have a hyperoceanic to oceanic climate
and a distinctive vegetation resulting from species of both Asian and North American origin.
The region supports shrublands, dwarf shrub heaths, and herbaceous meadows, with a flora
dominated by circumpolar Arctic and Asian and North American boreal species and a strong
component of more restricted amphi-Beringian species. But there are few species that are
endemic to this region or have only a North American distribution. While the bioclimate and
distinctive vegetation form the basis for a separate subdivision for this area, we included this
non-forested region in the Alaska–Yukon subdivision because the focus of this chapter is on
boreal forests.
13
Jean-Pierre Saucier, Ken Baldwin et al.
14
Boreal forests
and boreal plains. Associated ericaceous species are dominated by Ledum groenlandicum, Vaccinium
vitis-idaea and V. myrtilloides. Arctostaphylos uva-ursi, together with reindeer lichens (esp.
Cladonia2 mitis), is common on dry, poor sites. Alnus incana is largely associated with floodplains
in the West-Central region.
Distinction of the West–Central boreal from the Alaska–Yukon boreal corresponds to the
range limits of Pinus contorta var. latifolia and Abies lasiocarpa, as well as the increased prevalence
of northern species such as Vaccinium uliginosum, Empetrum nigrum, Betula nana, Arctous rubra and
Nephroma arcticum.
2 A recent decision about this genus put all Cladina species into Cladonia.We decided to use Cladonia even if Flora
of North America (1993+) still uses Cladina.
15
Jean-Pierre Saucier, Ken Baldwin et al.
stellaris and C. mitis cover the ground. On richer sites or warmer parts of the eastern boreal, the
herb layer is well developed to the detriment of the moss layer, while Acer spicatum, Corylus
cornuta and other broadleaved shrubs replace the heaths in the shrub layer. On wetter sites,
Sphagnum species, as well as shrub species such as Alnus incana ssp. rugosa, dominate the
understory.
16
Boreal forests
coast of Kamchatka, in an oceanic climate of mild winters, cool summers and a very short
growing season. About half of this area is in the zone of continuous permafrost. Compared to
other parts of the boreal zone, the Asian portion is the most diverse.
Altai–Sayan boreal
Southern Siberia is a region where the boreal zone transitions with temperate and steppe
regions. The Altai–Sayan floristic subdivision is influenced by the vegetation of adjacent
mountainous areas, including the major mountain systems from the Altai in the west to the
Hamar-Daban in the east.
Zonal vegetation on the low elevation arid plains is grass steppe vegetation, however in the
mountains, mixedwood (Betula pendula, Populus tremula, Pinus sylvestris) and conifer forests
(Pinus sibirica, Abies sibirica) prevail. These mountain systems are affected by humid Atlantic air
masses and receive over 1500 mm of annual precipitation. The conditions of relatively high
temperatures and high humidity that correspond to the temperate zone in central Eurasia have
led to the formation of unique forests characterized by boreal dominants and well-developed
layers of shrubs and tall herbs. These forests have a complete set of boreal species but differ from
other boreal forests by containing a very high proportion of temperate species in the understory
(Nazimova et al. 2014).
17
Jean-Pierre Saucier, Ken Baldwin et al.
Transbaikalian boreal
This floristic subdivision lies south of Lake Baikal and spreads from the Northern Baikal Plateau
in the west to the upper part of the Amur River in the east. The southern boundary of the
Transbaikalian boreal is in northeastern Mongolia. The climate is ultracontinental and, although
continuous permafrost is absent, soils are characterized by severe freezing. The landscape is
semi-forested, moisture being the major limiting factor for forest distribution. Forests mainly
occupy the northern aspects of mountain slopes, while southern aspects and valleys are covered
by steppe vegetation. Most Transbaikalian boreal forests are composed of Larix gmelinii, except
on sandy sites, which are occupied by Pinus sylvestris. At higher elevations, Pinus sibirica can
form sparse stands.
Typical Larix gmelinii forests occur within the forest–steppe ecotone of the Transbaikalian
boreal. Their open canopy is dominated by Larix gmelinii and Betula platyphylla in different
18
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Talks with Tolstoi
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Translator: S. S. Koteliansky
Virginia Woolf
Language: English
Credits: E-text prepared by Tim Lindell, Martin Pettit, and the Online
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BY
A. B. GOLDENVEIZER
TRANSLATED BY
S. S. KOTELIANSKY AND VIRGINIA WOOLF
August 2nd, 4 P.M. I have just had a long talk with L. N. on art. He
was repeating the contents of his article on art which he is writing,
and which he goes on working over and rewriting. In the course of it
L. N. said:
“When art became the inheritance of a small circle of rich people,
and left its main course, it entered the cul-de-sac in which we see it
now.
“Art is the expression of feeling, and the higher it is the greater the
public which it can draw to itself. Therefore the highest art must
reflect those states of mind which are religious in the best sense of
the word, as they are the most universal and typical of all human
beings.
“The majority of so-called works of art consist in a more or less
skilful combination of four elements: (1) borrowing—for instance,
the working out of some legend in a poem, of a song in music, etc.
Or unconscious borrowing—that is, an imitation now of one thing,
now of another, not intended by the author. (2) Embellishments:
pretty metaphors which cover up insignificant ideas, flourishes in
music, ornament in architecture, etc. (3) Effects: violent colours in
painting, accumulated dissonances, sharp crescendos in music, and
so on. Finally, (4) the interest—that is, the desire to surprise by the
novelty of the method, by the new combination of colours, etc.
Modern works of art are usually distinguished by these four qualities.
“The following are the chief obstacles which hinder even very
remarkable men from creating true works of art: first,
professionalism—that is, a man ceases to be a man, but becomes a
poet, a painter, and does nothing but write books, compose music,
or paint pictures; wastes his gift on trifles and loses the power of
judging his work critically. The second, also a very serious obstacle,
is the school. You can’t teach art, as you cannot teach a man to be a
saint. True art is always original and new, and has no need of
preconceived models. The third obstacle, finally, is criticism, which,
as some one has justly said, is made up of fools’ ideas about wise
men.
“I know that my article will be received by most people as a series of
paradoxes, but I am convinced that I am right.”
L. N. is evidently much carried away by his work.
August 2nd. I have been here from July 27th (in Yasnaya Polyana).
A queer young man, K., came to L. N., and, on my asking him what
he was doing, he said that “he was the free son of air.” K. told L. N.
that he wanted to settle down in the country among the people.
L. N. in recounting it said:
“Of course, I did not advise him to do it. Usually nothing comes from
such attempts. For instance, some very nice people, the N. N.’s,
bought a small plot of land and settled like that in the country. A
peasant cut down one of their trees; they did not want to take
action in the court against him, and soon, when the peasants learnt
about it, they cut down the whole woods. The peasant boys stole
their peas; they were not beaten nor driven away, and then nearly
the whole village came and stole all the peas, etc., etc.
“One should not, above all, look for new ways of life, for usually, in
doing so, one’s whole energy is spent on the external arrangement
of life. And when all the external arrangement is over, one begins to
feel bored and does nothing. Let every one first do his own work, if
only it does not clash sharply with his convictions, and let him try to
become better and better in his own situation, and then he will find
new ways of life into the bargain. For the most part, all the external
side of life must be neglected; one should not bother about it. Do
your own work.”
To-day L. N. said of some one:
“He is a Tolstoian—that is, a man with convictions utterly opposed to
mine.”
Yesterday L. N. spoke of the process of creative work:
“I can’t understand how any one can write without rewriting
everything over and over again. I scarcely ever re-read my published
writings, but if by chance I come across a page, it always strikes me:
All this must be rewritten; this is how I should have written it....
“I am always interested to trace the moment, which comes quite
early, when the public is satisfied; and the artist thinks: They say it
is good; but it is just at this point that the real work begins!”
To-day L. N. was not well. I went to him; he was lying on the little
sofa in the drawing-room. He told me of S. G. Verus’s book on the
Gospels.
“His final conclusion is the denial of Christ as a historical person. In
the earliest written parts of the New Testament—in Paul’s messages
—there is not a single biographical fact about Christ. All the Gospels
that have come down to us were composed between the second and
fourth century A.D. Of the writers who were Christ’s contemporaries
(Tacitus, Suetonius, Philo, J. Flavius) not a single one of them
mentions Christ; so that his personality is not historical, but
legendary.
“All this is very interesting and even valuable, for it makes it
unnecessary to quarrel any more over refuting the authenticity of
the Gospel stories about the miracles; and it proves the teaching of
the Gospels to be not the words of one superman, but the sum of
the wisdom of all the best moral teaching expressed by many people
and at different times.”
L. N. also said this to me:
“Perhaps it is because I am unwell, but at moments to-day I am
simply driven to despair by everything that is going on in the world:
the new form of oath, the revolting proclamation about enlisting
university students in the army, the Dreyfus affair, the situation in
Serbia, the horrors of the diseases and deaths in the Auerbach
quicksilver works.... I can’t make out how mankind can go on living
like this, with the sight of all this horror round them!
“It always strikes me how little man is valued, even in the simplest
way as a valuable and useful animal. We value a horse which can
carry, but man can also make boots, work in a factory, play the
piano! And 50 per cent are dying! When I used to breed merino
sheep and their death-rate reached 5 per cent, I was indignant and
thought the shepherd very bad. And 50 per cent of the people are
dying!”
I read L. N.’s most wonderful Father Sergius.
Moscow, August 9th. I returned from Yasnaya in the evening of the
6th. This is what I find I have written down.
The talk turned upon the woman question. The conversation was
carried on in a half-jocular tone.
L. N. said:
“Woman, as a Christian, has a right to equality. Woman, as member
of the modern and perfectly pagan family, must not struggle for an
impossible equality. The modern family is like a tiny little boat sailing
in a storm on the vast ocean. It can keep afloat if it is ruled by one
will. But when those in the boat begin struggling, the boat is upset,
and the result is what we see now in most families. The man,
however bad, is in the majority of cases the more sensible of the
two. Woman is nearly always in opposition to any progress. When
man wants to break with the old life and to go ahead, he nearly
always meets with energetic resistance from the woman. The wife
catches hold of his coat-tails and will not allow him. In woman a
great evil is terribly highly developed—family egotism. It is a
dreadful egotism, for it commits the greatest cruelties in the name of
love; as if to say, let the whole world perish so that my Serge may
be happy!...”
Then L. N. recalled scenes which he had observed in Moscow:
“There issues from Minangua’s a gentleman in a beaver coat, with a
sad face, and after him his lady, and the porter carries boxes and
helps the lady into the sledge.
“I love at times to stand near the colonnade by the great theatre
and watch the ladies driving up to stop at Meriliz’s. I only know of
two similar sights: (1) when peasant women go to Zaseka to pick up
nuts the watchmen catch them, so that sometimes they give birth
out of fright, and yet they go on doing it; and (2) so it is with ladies
shopping at sales.
“And their coachmen wait in the bitter cold and talk among
themselves: ‘My lady must have spent five thousand to-day!’
“I shall one day write about women. When I am quite old, and my
digestion is completely out of order, and I am still looking out into
the world through one eye, then I shall pop my head out and tell
them: That’s what you are! and disappear completely, or they would
peck me to death.” ...
Doctor E. N. Maliutin was in Yasnaya. L. N. said to him:
“I can’t understand the usual attitude that a doctor always serves a
good cause. There is no profession that is good in itself. One may be
a cobbler and be better and nicer than a doctor. Why is restoring
some one to health good? At times it is quite the opposite. Man’s
deeds are good, not in themselves, but because of the feelings
which inspire him. That’s why I do not understand the desire of
women to be doctors, trained nurses, midwives, as though by
becoming a midwife everything is settled for the best.”
On some occasion L. N. said:
“When you are told about a complicated and difficult affair, for the
most part about some one’s disgusting behaviour, reply to it: Did you
make the jam? or: Won’t you like to have tea?—and that’s all. Much
harm comes from the so-called attempt to understand circumstances
and relations.”
December 7th. When Tolstoi was ill (he is much better now) and I
was for the first time in his room, he seemed glad to see me, which
was a great delight to me. On his table was the volume of Tyutchev’s
poems. In his hand he had an English book, Empire and Freedom (I
don’t remember by whom). As is always his way, Tolstoi at once
spoke of what he was reading.
“Here is a remarkable book!” said Tolstoi. “He (the author) is
American, therefore an Anglo-Saxon; nevertheless, he denies the so-
called civilizing influence of the Anglo-Saxon race. I can’t understand
how people can stick to such superstitions! I understand a
Muhammad preaching his doctrine,—mediæval Christianity, the
Crusades. Whatever the convictions of those people may have been,
they did it in the belief that they knew the truth and were giving that
knowledge to others. But now there is nothing! Everything is done
for the sake of profit!”
Then Tolstoi began to talk about a French pamphlet on the workers’
co-operative societies which he had read.
“Why not introduce in the villages here such co-operative societies?
That is a vital thing! You, instead of doing nothing,” he turned to Ilya
Lvovich, who sat there, “ought to do it here in the village.
“Socialist ideas have become a truism. Who can now seriously
dispute the idea that every one should have the right to enjoy the
result of his labour?”
Then the conversation turned upon the obschina.
Tolstoi said:
“Everything is taken away from the peasants; they are overtaxed,
oppressed in all ways. The only good thing left is the obschina. And
then every one criticizes it and makes it responsible for all the
miseries of the peasants, in their wish to take away from the
peasants their last good thing. They make out that the mutual
responsibility of the members is one of the evils of the obschina. But
mutual responsibility is only one of the principles of the obschina
with regard to fiscal purposes. If I use a good thing for an evil end,
that does not prove that the thing is in itself bad.”
Then the conversation turned upon Tyutchev. The other day Tolstoi
saw in the Novoe Vremya his poem “Twilight.” He therefore took
down all Tyutchev’s poems and read them during his illness.
Tolstoi said to me:
“I am always saying that a work of art is either so good that there is
no standard by which to define its qualities—that is real art,—or it is
quite bad. Now, I am happy to have found a real work of art. I
cannot read it without tears. I know it by heart. Listen, I’ll read it to
you.”
Tolstoi began in a voice broken with tears:
“The dove-coloured shadows melted together....”
When I am on my death-bed I shall not forget the impression then
produced on me by Tolstoi. He lay on his back, convulsively twisting
the edge of his blanket with his fingers and trying in vain to restrain
the tears that choked him. He broke down several times and began
again. But at last, when he read the end of the stanza, “Everything
is in me, and I in everything,” his voice gave way. The entrance of A.
N. Dunaev stopped him. He grew calmer.
“What a pity that I spoilt the poem for you!” he said to me later.
Then I played the piano.
Tolstoi asked me not to play Chopin, saying: “I am afraid I might
burst into tears.”
Tolstoi asked for something by Mozart or Haydn.
He asked: “Why do pianists never play Haydn? You ought to. How
good it is—beside a modern complicated, artificial work—to play
something of Mozart or Haydn!”