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Forest Ecology and Management 259 (2010) 2172–2182
Contents lists available at ScienceDirect
Forest Ecology and Management
journal homepage: www.elsevier.com/locate/foreco
Beech regeneration research: From ecological to silvicultural aspects
Sven Wagner a,∗ , Catherine Collet b,c , Palle Madsen d , Tohru Nakashizuka e ,
Ralph D. Nyland f , Khosro Sagheb-Talebi g
a
TU-Dresden, 01735 Tharandt, Germany
AgroParisTech, UMR1092, Laboratoire d Etude des Ressources, Foret Bois (LERFoB), ENGREF, 14 rue Girardet, 54000 Nancy, France
c
INRA, UMR1092, Laboratoire d Etude des Ressources, Foret Bois (LERFoB), Centre INRA de Nancy, 54280 Champenoux, France
d
Forest and Landscape Denmark, Vejle, Denmark
e
Graduate School of Life Sciences, Tohoku University, 6-3 Aoba, Aramaki, Aoba-ku, Sendai 980-8578, Japan
f
SUNY College of Environmental Science and Forestry, Syracuse, NY, USA
g
Research Institute of Forests and Rangelands, Tehran, Iran
b
a r t i c l e
i n f o
Article history:
Received 2 March 2009
Received in revised form 12 February 2010
Accepted 28 February 2010
Keywords:
Fagus
Model-genus
Life cycle
Regeneration measures
a b s t r a c t
This review describes key regeneration characteristics of the genus Fagus as represented by its four most
prominent species: F. crenata (F.c.), F. grandifolia (F.g.), F. orientalis (F.o.) and F. sylvatica (F.s.). Similarities
and differences in the relevant life phases of these species are identified. Those are related to natural
disturbance regimes and synecological peculiarities of the forests where they grow, thereby establishing
a basis for evaluating the likely outcome of different silvicultural measures.
Common ecological characteristics of these Fagus species’ life cycles include the masting phenomenon,
pollen dispersal with effective distances of about 100 m, seed dispersal to about 20 m, seedling sensitivity
to frost, drought, and animal predation, and a very shade tolerant establishment phase. This commonality
suggests its appropriateness as a “model-genus”. However, some species also have unique ecological
characteristics not observed in the others. F.g. exhibits root suckering, and beech bark disease seems
to trigger vegetative regeneration by that means. Likewise, its masting behaviour deviates from F.s. F.o.
and F.c., occurring more frequently and more regularly. In F.c. forests, dwarf bamboo species and their
ecological characteristics are important determinants of tree regeneration establishment.
The small canopy gaps that commonly occur in Fagus dominated natural forests fit very well with
the genus’ regeneration characteristics. These conditions are best duplicated by management measures,
which maintain partial overstory shading until the seedlings are large enough for release. However,
such a strategy reduces chances to regenerate more light-demanding associated species. Together with
differences in landowner objectives, the diversity of ecological conditions within and between the species
of Fagus requires site-specific prescriptions to insure regeneration success, e.g. cutting regimes.
Of particular interest to research are the challenges of managing mixed-species stands for high quality
timber production in Central European and Caspian beech forests, the decline of F.g. and how to deal with
the aftermath forest, and effective ways to manage F.c. in coexistence with dwarf bamboo. Further, the
historic dispersal of heavy seeded Fagus species over long distances is still poorly understood. In addition,
since their drought sensitive seedlings may be damaged or killed during extreme weather, research must
address the possible effects of global climate change on the regeneration potential of beech forests.
Species-bridging research may be needed to address these questions.
© 2010 Elsevier B.V. All rights reserved.
1. Introduction
Pressure to apply ecologically sound practices and make decisions more transparent requires forest scientists to explore new
pathways for managing forest stands. Global dialogue provides
∗ Corresponding author. Tel.: +49 35203 3831300; fax: +49 35203 3831397.
E-mail address: wagner@forst.tu-dresden.de (S. Wagner).
0378-1127/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2010.02.029
ideas and solutions. But silviculture must ultimately account for
regional differences (e.g., species characteristics and mixtures,
environment, and management objectives) that affect the outcome
of alternative treatments. As a contribution, we introduce the concept of a “model genus” using the species of Fagus as an example.
A similarity in their features allows the integration of information from multiple, independent studies, and provides a basis for
evaluating alternative management strategies for beech dominated
forests.
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Among features that make Fagus suitable to this approach are:
- its distribution throughout the temperate northern hemisphere
(Peters, 1997), and the range of associated environmental conditions;
- the ecological similarity of its species as representatives of ones
having shade-tolerant, dominant climax traits;
- the amount and quality of available knowledge about its biology
and management; and
- the economical importance of its component species.
This review makes that vision real by synthesizing detailed autand synecology research findings about Fagus regeneration, and
articulating the similarities and differences in ecologic characteristics and silviculture for the genus around the world. It primarily
compares Fagus crenata Blume (F.c.), Fagus grandifolia Ehrh. (F.g.),
Fragus orientalis Lipsky (F.o.) and Fagus sylvatica L. (F.s.). Oriental
beech (F.o.) is included as a species, as it has its own morphological
trait. We are aware of the ongoing discussion about the species status of F.o. as it has few unique alleles (see Denk et al., 2002; Salehi
Shanjani and Sagheb-Talebi, 2006; Paffetti et al., 2007). Only limited information about other Fagus species is available, which were
not taken into account in the review.
We structured the review around the different life cycle phases
of these species (after Harper, 1977), including:
- flowering and fruiting phase
- seed dispersal phase
- establishment phase
By comparing both differences and similarities among the
species of Fagus, we identified the risks of failure in management
measures and the means to overcome them. Such an approach
insures that silviculture accounts for the critical ecologic factors
and results in an acceptable vitality, density, survival, growth, and
quality of regeneration at an appropriate time.
The review: (i) tests the applicability of the model genus idea
using Fagus; (ii) identifies regeneration problems with different
Fagus species; (iii) discusses options for reducing the risks; and (iv)
identifies research needs.
2. Beech regeneration in the context of forest dynamics
2.1. Pattern of forest disturbance, natural regeneration and the
silvicultural implications
The majority of North American and European beech forests
have been logged or once cleared for agricultural use (Peters,
1997), followed by replenishment or recolonization through natural regeneration. In fact, primary forests having an important
component of beech are absent or extremely rare on these two
continents (Cronon, 1983; Bradshaw, “Holocene history of beech
Fig. 1. Canopy gap size frequency distributions of four beech forest reserves.
Log-normal distributions were computed for two F.c. forests (data published in
Nakashizuka, 1984; Yamamoto, 1989) and two F.s. forests (data published in Tabaku
and Meyer, 1999; Zeibig et al., 2005) by maximum-likelihood method.
forests”, this volume). However, Japan still has about 1.4 million ha
of primary forest (Ministry of Environment, 1997) and Iran more
than 100,000 ha with F.o.as an important species (Sagheb-Talebi
et al., 2004; Knapp, 2005). These remaining areas provide a valuable reference for studying temperate deciduous forest ecosystem
dynamics, and when considering the silvicultural options for beech
forests in general.
Forest disturbance is central to regeneration responses of Fagus
throughout the globe, both in primary forests and replenished or
managed ones. In that sense, “disturbance” includes anything that
initiates regeneration from seeds, affects the growth dynamics of
advance seedlings and saplings, or induces root suckering or stump
sprouting.
2.1.1. Pattern of forest disturbance
Disturbance in beech forests may range from large-scale standreplacing events (blowdown during a storm), to large openings
once occupied by a group of trees, to small single tree gaps. Mortality of small or intermediate trees due to inter-tree competition
also open limited-size gaps in the canopy (Nakashizuka et al., 1992;
Delfan Abazari et al., 2004a,b). These disturbances may alter the
forest structure without reducing canopy closure, as with the dominance and synchronous death of dwarf bamboos (Sasa spp.) in
F.c. ecosystems (Abe et al., 2001), or when disturbance to shallow
roots triggers root suckering of F.g. (Nyland et al., 2006a). Prolonged
browsing during times of high deer densities has also altered the
composition and abundance of tree seedlings and other vegetation
in F.s. and F.g. stands (e.g. Sage et al., 2003). Air pollution, nutrient deposition, and altered ground water tables may also affect
forest structure and regeneration. Even so, “disturbance” in temperate regions is commonly equated with openings (gaps) in the
overstory canopy. Further, the multi-age and irregular structure
Table 1
Indicating variables of canopy gap size frequency distributions of some beech forest reserves. Log-normal distributions were computed by maximum-likelihood method.
Values of indicating variables were taken from those computations.
Species, authors,
name of reserve,
area considered
F. crenata, Nakashizuka, 1984, Mt. Moriyoshi, 2.4 ha
F. crenata, Yamamoto, 1989 Miscellaneous, 20.0 ha
F. sylvatica, Zeibig et al., 2005, Krokar, 12.0 ha
F. sylvatica, Tabaku and Meyer, 1999, Mirdita, 5.0 ha
F. sylvatica, Tabaku and Meyer, 1999, Puka, 3.6 ha
F. sylvatica, Tabaku and Meyer, 1999, Rajka, 6.0 ha
Average gap size
(m2 )
161
59
129
74
62
67
Mode of gap size
(m2 )
Proportion of total
gap area in gaps
smaller than
200 m2
Proportion of
reserve area under
gaps, gap definition
39
34
43
39
46
50
0.40
0.95
0.53
0.88
0.99
0.99
0.20 canopy gaps
0.12 canopy gaps
0.056 canopy gaps
0.066 canopy gaps
0.034 canopy gaps
0.033 canopy gaps
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of primary F.o. stands of northern Iran (Eslami and Sagheb-Talebi,
2007; Shahnavazi et al., 2005; Sagheb-Talebi and Schütz, 2002) suggests that the patterns of disturbance and regeneration may differ
among developmental stages and stand types within a region.
2.1.1.1. Gap size distribution. Canopy disturbance promotes the
growth of beech regeneration into upper canopy positions. Thus,
the frequency and sizes of gaps profoundly affects the dynamics
of beech communities. In turn, the species composition of a stand
and local environmental conditions temper the disturbance regime.
However, within pure beech forests, gaps are primarily small (F.c.:
Nakashizuka, 1987; Yamamoto, 1989; F.g.: Runkle, 1982; F.s.: Butler
Manning, 2007).
Fig. 1 and Table 1 describe gap frequencies for different forests
having pure beech stands. They show that gaps smaller than 200 m2
are frequent in old-growth beech-dominated systems (mean gap
size of 59 to 161 m2 ). With F.c. and F.s. most gaps are between 34 and
50 m2 , similar to the crown size of old beech trees and suggesting
a single tree mortality pattern of gap formation. While naturally
created forest gaps in most beech forests may cover between 3 and
20% of the area (Table 1), Japanese forests have 12–20% in gaps
(Nakashizuka, 1987).
2.1.1.2. Disturbance caused by beech bark disease. Understory F.g.
density has increased as beech bark disease spread across northeastern North America following the introduction of Cryptococcus
fagisuga (Lindinger) from Europe. The scale insect creates openings in the bark. These serve as infection courts for a Nectria fungus
that kill patches of cambium, eventually girdling the tree (Houston,
2005). Since only about 1% of all F.g. may have a resistance, tolerance, or immunity (Houston, 1997), few trees larger than 20–25 cm
diameter escape its effect (Houston, 1997). Similar symptoms have
recently been reported in Caspian beech forests (Kiadaliri et al.,
2008).
Where F.g. has been abundant in a stand, beech bark disease
opened the overstory canopy appreciably, and root suckers have
become dense throughout the understory (Nyland et al., 2006a).
Within stands having only scattered F.g. trees, the mortality creates
dispersed small gaps, and root suckers mostly encircle the dying
trees.
2.1.1.3. Disturbance within stands having a dwarf bamboo understory. Survival and development of F.c. regeneration depends on
light availability, and this is influenced by the amount of dwarf
bamboo biomass in an understory (Maeda, 1988; Abe et al., 2001,
2002). The dwarf bamboo flowers once in several decades and then
dies over even several square kilometres, dramatically changing
the understory light regime (Nakashizuka, 1987; Abe et al., 2001)
and stimulating growth of any advance beech regeneration. Hence,
synchronous canopy gap formation and bamboo dieback seem necessary for beech regeneration to become established and grow.
2.1.2. Sexual regeneration
2.1.2.1. Flowering, pollen dispersal, and fruiting. Male and female
flowers of beech occur separately on the same tree, and are vulnerable to spring frosts (Savill, 1991; Young and Young, 1992). F.s.
pollen effectively disperses less than 250 m within forests (Wang,
2001; Gerber in Kramer, 2004), and F.c. up to 193 m (Shimatani et al.,
2007). Pollen may disperse for 500 m in some cases, but generally
only up to 60 m in high density stands (Shimatani et al., 2007).
Evidence relates seed production to tree diameter (for F.s.:
Wagner, 1999), with flowering and seed production beginning at
about age of 40 to 50 in F.g., F.c., and F.s. Major masting events in
the later two species occur at 2- to 20-year intervals (Watt, 1923;
Young and Young, 1992; Kon et al., 2005). F.o. generally begins fruiting at age 60 with masting every 3 to 18 years (Mirbadin and Gorji,
Fig. 2. Cumulative beech dispersal models derived from established seedlings distributions. Models of Shimatani et al. (2007) for F.c., and Gerber (in Kramer, 2004)
for F.s. based on genetic analysis in pure stands. Model by Irmscher (unpubl. data)
for F.s. based on the distribution of established F.s. seedlings from a single mother
tree in a pure spruce (Picea abies) stand.
1996; Mirbadin and Namiranian, 2005). By contrast, intervals of
2-3 years seem common for F.g. (Schopmeyer, 1974; Jacabus et
al., 2005; McNulty and Masters, 2005). And while seed production
among trees >25–30 cm dbh decreased by two-thirds to threefifths as beech bark disease progressed (Costello, 1992), masting
recently increased due to modest seed production on large numbers of smaller trees that eventually reached pole stage (McNulty
and Masters, 2005).
Good seed production in F.s. and F.g. depends on climatic events
during two consecutive years having conditions that favour carbohydrate build-up, followed by an early summer drought the
next year (Piovesan and Adams, 2001). A warm July portends good
flowering of F.s. the following year (Övergaard et al., 2007), and
frequency of masting has positively correlated with site index. F.o.
flowering and fruiting also depend on climate and site conditions,
with heavy seed production correlated with soil nutrition, northern aspects, and an altitude of around 1500 m.a.s.l. Seeds also had
greater size and weight in stands at 750 to 1500 m.a.s.l. (Etemad
and Marvi Mohajer, 2004). Fruiting of F.o. in the Caspian region
varies from tree to tree, and site to site. That, along with the frequent masting of F.g., seems contradictory to the hypothesis for a
climatic linkage to beech masting, and consistent with observations
by Masaki et al. (2008) that masting events in F.c. differ among the
districts in northern Japan.
Theory predicts an evolutionary benefit for masting, particularly
with respect to pre-dispersal seed predation (Sork, 1993; Yasaka
et al., 2003; Piovesan and Adams, 2005). Different factors seem to
control these events with F.s. and F.c., compared to F.g. (Piovesan
and Adams, 2001), but they are poorly understood. This problem
is much more pressing in Japan, where managers need to predict the timing of good seed years for F.c. so they can schedule
advance clearing of dwarf bamboo from the understory. Yet after
considering several hypotheses forwarded by Kelly and Sork (2002)
and Kelly (2004), Yasaka et al. (2003) and Kon et al. (2005) concluded that escape from predation and pollination efficiency are
most important determinants for F.c.
2.1.2.2. Dispersal. Beechnuts commonly disperse by barochory,
usually for up to 20 m. Yet more than 80 m has been observed with
F.c. and F.s. Models of established beech regeneration for F.c. by
Shimatani et al. (2007) and F.s. by Gerber (in Kramer, 2004) appear
in Fig. 2, along with one by Irmscher (unpubl. results). They suggest
distances of up to 125 m, with zoochorous dispersal even introducing beech into stands of other species.
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S. Wagner et al. / Forest Ecology and Management 259 (2010) 2172–2182
Small mammals generally move nuts of F.c. and F.g., but for relatively short distances (Tubbs and Houston, 1990; Young and Young,
1992; Miguchi, 1996). Yet Kunstler et al. (2004) found F.s. natural regeneration in pine stands up to 3,000 m from nearest beech
trees, compared to less than 300 m into open grassland. Jays seemed
primarily responsible for the longer dispersal. The birds also may
carry F.g. seeds for several kilometres (Johnson and Adkisson, 1985;
Tubbs and Houston, 1990).
2.1.2.3. Seed wintering and germination. In F.g., burs open in
autumn after freezing temperatures, commonly releasing two nuts
per bur (Burns and Honkala, 1990). The seeds fall to the forest floor
in a dormant state and remain so throughout autumn and winter. F.s. nuts need several months of this pre-chilling, depending
on the temperatures during that period (Tubbs and Houston, 1990;
Gosling, 1991; Young and Young, 1992). Individual seeds and different seedlots vary in these requirements, but natural stratification
in a moist seedbed usually pre-conditions them. Since germination and sprouting are temperature dependent (Harper, 1977), the
time of germination varies with spring weather conditions (Runkle,
1989). Radicles may emerge several months before the embryonic
shoot develops into a recognizable beech seedling (Madsen, 1995;
Farmer, 1997; Young and Young, 1992).
The requisite pre-chilling complicates programs of artificial
regeneration with Fagus (i.e., nursery production or direct seeding),
since managers depend on reliable and predictable germination following sowing (Willoughby et al., 2004a; Baumhauer et al., 2005).
Bonner and Leak (2008) described techniques for pre-chilling an
entire beech seedlot at 28–30% moisture content until dormancy
is broken in most of the seed, then increasing moisture to initiate
uniform germination.
Beechnuts landing on natural seedbeds are susceptible to
unfavourable environmental conditions (e.g. desiccation and frost),
harmful fungi, and predation (insects, birds, and both large and
small mammals). The most important predators of F.s. are rodents
such as Apodemus sylvatica, Clethrionomys glareolus, birds and wild
boar (Sus scrofa) (Burschel et al., 1964; Harmer, 1995; Ammer et
al., 2002; Willoughby et al., 2004b). Survival has been improved
dramatically by seedbed preparation shortly before, during, or
after natural seed fall or direct seeding (See Burschel et al., 1964;
Huss and Burschel, 1972; Huss and Stephani, 1978; Jungbluth and
Dimitri, 1980; Madsen, 1995). For direct seeding, Ammer et al.
(2002) demonstrated that mineral soil seedbeds provide a stable
moisture regime and soil temperatures favourable to beechnut
germination. In fact, first-year regeneration density may be 100fold greater in prepared seedbeds than on the untreated forest
floor (Olesen and Madsen, 2008). Liming acidic soil also improved
seedling density and growth following direct seeding (Küssner and
Wickel, 1998; Ammer et al., 2002), but had no effect at less acidic
sites.
Studies of mixed soil and mineral seedbeds did not reveal
whether lower regeneration density in the former may result from
increased fungal attack due to the higher organic matter content in
the soil, or because the mixed seedbed facilitated rodent tunnelling
and seed predation (Burschel et al., 1964; Dubbel, 1989; Madsen,
1995). Snow cover has reduced predation of F.c. nuts (Shimano and
Masuzawa, 1998; Homma et al., 1999).
Many birds, mammals, and insects eat or damage F.g. and F.s.
seeds, and they affect dispersal by collecting and storing beechnuts in caches. Beechnut availability may affect the survival and
reproduction of these creatures during harsh winters (Jensen, 1985;
Yasaka et al., 2003; Jacabus et al., 2005), with rodent populations increasing following major beech masting events (Jensen,
1982). Calculations of potential consumption based on energy
requirements suggest that small animals may consume significant
proportions of the beechnut crop during years of limited masting.
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However, during major masts years the loss to all kinds of predation
will have little effect on overall seed supply (Jensen, 1982; Yasaka
et al., 2003; Kon et al., 2005; Olesen and Madsen, 2008).
2.1.2.4. Establishment and early growth.
2.1.2.4.1. Effects of different canopy openings. Beech seedlings
become established under a wide range of canopy openings
(Sagheb-Talebi and Schütz, 2002). They survive for long periods
at very low light levels (Relative Light Intensity, RLI = 1%) (Emborg,
1998; Modry et al., 2004), but grow slowly (Gansert and Sprick,
1998; Collet et al., 2001). Height and diameter growth are best in
the open (RLI = 100%), but differ little with light at 30% < RLI < 50%
(Gemmel et al., 1996; Kunstler et al., 2005). Beech seedlings often
undergo multiple suppression-and-release episodes before reaching the upper canopy (Nagel et al., 2006; Collet et al., 2008). Even
after a long period of suppression, height growth increases following each canopy disturbance, (Nakashizuka, 1983; Canham, 1990;
Collet and Chénost, 2006) and particularly after the second and
third release (Leak, 2003).
Seedlings have a highly plastic morphology that depends on
genetics, light, water, nutrient availability, and frost occurrence
(Thiébaut et al., 1985; Nicolini, 1997). Beech is characterized by
a monopodial branching pattern, a plagiotropic trunk secondarily reoriented into a vertical position by cambial activity, and
plagiotropic branches (Peters, 1997; Hallé et al., 1978). Shoot elongation is rhythmic, with two growth flushes per season at good
sites and often sympodial due to frequent death of the apical bud
(Sagheb-Talebi, 1996; Roloff, 1986). At RLI > 40%, all main axes orient into a vertical position, but multiple growth flushes and the
rapid growth induced by high light result in large branches and
forks. With RLI < 10%, the branches and main axis of beech do not
reach a vertical position due to the low radial growth (Nicolini et
al., 2001), negatively affecting sapling architecture (for F.s.: Diaci
and Kozjek, 2005; for F.g.: Canham, 1988; Nyland et al., 2006a).
However, branches and forks remain small in diameter and do
not compete strongly with the main axis (Bonosi, 2006). At light
20% < RLI < 40%, F.s. and F.o. seedlings have a better morphology,
with weaker lateral branches and forks than at high light levels
and greater verticality of the main axis than at low light levels
(Sagheb-Talebi et al., 2001).
Overtopping mature trees and neighbouring understory vegetation may shade beech seedlings. Yet moderate to dense shading
over long periods may result in better-formed beech saplings
(Leonhardt and Wagner, 2006). At full light, beech will have good
morphology only when grown at a high seedling density of conspecifics (Sagheb-Talebi and Schütz, 2006) or with neighboring
vegetation of comparable density. To that end, recommendations
for low-density plantations (e.g. fewer than 1500 tree per ha)
include maintaining or establishing neighbouring woody vegetation to provide the necessary lateral shade.
2.1.2.4.2. Limitations in analysis of single factors. Most studies
of beech seedling responses have not separated effects of competition for various resources, complicating any extrapolation to sites
with different water or nutrient availabilities (but see Wagner et
al., 2009). Height growth at high light is much less at low than at
high water availability (Vinkler et al., 2007) and growth responses
after changes in light availability also depend on water availability (Madsen, 1994). Seedling growth has also been related to light
availability and root density of old beech (Wagner, 1999), with
F.s. seedlings recovering more quickly in good light after trenching excluded roots of older trees (Fig. 3). Overall, RLI seems the
best indicator of environmental change, with morphology of young
beech differing under varying light environments.
2.1.2.4.3. Ground vegetation as competitor. Dwarf bamboo prevents development of even advance F.c. regeneration (Abe et al.,
2002), and associated ground vegetation affects F.s. seedling growth
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S. Wagner et al. / Forest Ecology and Management 259 (2010) 2172–2182
Fig. 3. F.s. seedling growth related to Relative Light Intensity (RLI in percent of that
above canopy). Open circles indicate relative height growth prior to trenching in
a pure beech stand. Solid circles indicate relative height growth in the same plots
2 years after root trenching. The
are
based on the Michaelis-Menten
predictions
function RHG = A (RLI − B) /
A/C
+ (RLI − B)
, with RHG as relative height
growth, RLI as relative light intensity and A, B and C the parameters. Data from
Wagner (1999).
(e.g., Burschel and Schmaltz, 1965; von Lüpke, 1987). While beech
seedlings and saplings withstand interference by most weeds
(Madsen, 1995), they must be weeded to insure satisfactory height
growth when planted or sowed on former farmland (Löf, 2000; Löf
et al., 2004). Much depends on the density of ground vegetation
and the effect of any overstory trees.
Most ground vegetation does not thrive as well as young beech
beneath a moderately closed F.s. canopy, lessening its competitive
effects. In fact, site preparation using herbicides did not improve
F.s. seedlings biomass production when applied under a dense shelter, compared to a light overhead cover (Huss and Stephani, 1978),
suggesting use of moderate canopy openings to establish F.s. regeneration where competition from ground vegetation might be strong
(Kühne and Bartsch, 2003).
Compared to oaks, planted F.s. seedlings are more prone to mice
and vole damage in areas with weed cover (von Lüpke, 1987). With
F.c., predation by rodents beneath closed-canopy stands decreases
following the dieback of dwarf bamboo, resulting in increased seed
and seedling survival (Abe et al., 2005).
2.1.3. Vegetative regeneration
2.1.3.1. Sprouting and root suckers. While F.c., F.g., F.o. and F.s.
sprout from cut stumps, coppice systems have been developed only
for F.c. and F.s. (Peters, 1997). In F.g., stump sprouts originate either
from adventitious buds around the top of a stump, or from dormant
buds along the sides (Mallett, 2002; Nyland et al., 2006a). Most die
within 2–3 years (Mallett, 2002), but some persist off stumps in
the open or beneath a low-density overstory (Nyland, unpublished
results). Sprouting potential decreases after F.g. and F.s. trees reach
10 cm dbh, and most F.g. stump sprouts do not develop into trees
(Eyre and Zillgitt, 1953; Hamilton, 1955; Fowells, 1965).
F.g. produces root suckers. These may form a dense understory following canopy disturbances (Houston, 1975; Krasny and
DiGregorio, 2001; Nyland et al., 2006a; Runkle, 2007), usually
within 9–10 m around the source trees (Jones and Raynal, 1987;
Nyland et al., 2006a). Most arise from wounds to shallow and
exposed roots (Jones and Raynal, 1987), and often following logging (Houston, 1975; Jones and Raynal, 1987; Houston, 2001).
Yet dense root suckers have developed even beneath unmanaged
stands (Nyland et al., 2006a). As the suckers develop, new ones
originate off their extending roots, so that groups of overstory F.g.
trees may originate from a common source (Houston and Houston,
1987). As a result, a stand with many individual F.g. stems may have
relatively few separate clones.
Among advance F.g. regeneration <1.2 m tall, seedlings may
be more abundant than suckers at some sites, but not at others
(Nyland, 2008). Therefore, each F.g. clone may have a characteristic
flowering and suckering tendency. Even so, at sites having many
small F.g. seedlings, root suckers commonly dominate the advance
beech >0.6 m tall (Nyland, 2008).
Root suckers have become dense in unmanaged stands where
beech bark disease weakened or killed the large F.g. (Nyland
et al., 2006b), including unmanaged old-growth communities
(McNulty and Masters, 2005). This implicates the disease as a probable contributing agent. Surface disturbance and light near the
ground seem linked to root suckering in managed stands infected
with beech bark disease (Nyland et al., 2006a). Yet root suckers
may be more abundant around resistant beech trees and their
stumps, than around diseased trees and their stumps (Houston,
2001).
2.2. Competition and predation in the establishment
phase—impact to mixtures
2.2.1. Height growth dynamics and interspecific competition
In the young stages, F.g. grows slower than Acer saccharum,
Betula alleghaniensis, or Fraxinus americana (Nyland et al., 2006a).
So does F.s. growing in mixture with Fraxinus excelsior, Acer pseudoplatanus, Quercus petraea, and Q. robur (Joyce et al., 1998; Hein et
al., 2008), and F.o. intermixed with Acer velutinum (Sagheb-Talebi,
1998). Yet Fagus seedlings overtopped by other species generally
persist for long periods, then regain and maintain dominance at
later stages of stand development (for F.c.: Yoshida and Kamitani,
2000). Thus, it is common to grow F.s. in mixture with other species,
waiting for it to eventually become dominant (Joyce et al., 1998).
This transition usually occurs early in a rotation for mixtures of F.s.
and small-stature pioneers (e.g., Betula pendula) or slowly growing
species (like Quercus petraea, Sorbus torminalis), but late in a rotation for stands having high-stature post-pioneer ones (e.g., Fraxinus
excelsior, Acer pseudoplatanus).
In mixed-species regeneration, the relative dominance of each
varies along trophic and water gradients (Ellenberg, 1988b). On
highly productive sites with rapidly growing post-pioneer species
(e.g., Fraxinus excelsior, Acer pseudoplatanus), F.s. seedlings may
succumb within a few years (Rysavy, 1991), whereas an opposite
dynamics is observed on nutrient-poor sites. Similarly, with FagusQuercus mixed regeneration, Quercus will dominate only on drier
sites (Vera, 2000).
The balance among species also depends on overstory and
ground vegetation competition, herbivory, effects of pathogens,
interactions with microfauna and microflora, and other abiotic
factors such as late frost (Connell, 1990). Research has mainly concentrated on overstory competition and ungulate herbivory. Both
strongly affect development of Fagus and its associated species, and
both may be controlled by well designed management practices.
2.2.2. Light availability and canopy disturbance
While Fagus seedlings generally grow more slowly than most
associated species (Beaudet and Messier, 1998; Dreyer et al., 2005),
they survive better at low or intermediate light (F.g., Logan, 1973;
Nyland et al., 2006a,b). Thus, managers can influence the composition of mixed-species regeneration by regulating the degree
of canopy cover (Abe et al., 1995; Poulson and Platt, 1996). To
illustrate, with mixtures of shade-intermediate Fraxinus excelsior
and Acer pseudoplatanus, maintaining a closed canopy reduces
their growth compared to F.s. With less shade-tolerant species
like Quercus petraea, an RLI < 30% will preferentially enhance the
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development of Fagus (von Lüpke, 1998; Shahnavazi et al., 2005;
Sagheb-Talebi et al., 2001).
Height growth of Fagus increases after release by natural disturbance (including beech bark disease) or overstory cutting, even
after long periods beneath a closed canopy (Canham, 1990; Collet
et al., 2001; Nyland et al., 2006b; Nagel et al., 2006; Emborg,
2007) . Given periodic release, the beech develops into tall advance
regeneration that may interfere with other species (Krasny and
DiGregorio, 2001; Collet et al., 2008). In fact, stands having a dense
understory of F.g. will lack other regeneration, necessitating beech
removal to insure establishment of other species (Kelty and Nyland,
1981; Bohn and Nyland, 2003; Hane, 2003).
2.2.3. Browsing
Beech seedlings are vulnerable to birds, rodents, deer, and
other herbivores (Burschel et al., 1964; Madsen, 1995; Olesen and
Madsen, 2008). Deer prefer Acer saccharum, Acer rubrum, Betula
alleghaniensis, Prunus serotina, Fraximus Americana, Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Quercus petraea and Q.
robur over F.g. and F.s (Harmer, 2001; Gill, 1992; Ellenberg, 1988a;
Eiberle and Bucher, 1989; Nyland et al., 2006a). They also feed
on succulent F.g. stump sprouts and root suckers (Nyland et al.,
2006a), but protracted browsing of other species commonly promotes the dominance of understory F.g., even beneath uncut stands
(Nyland et al., 2006a). In F.c. forests, browsing reduces dwarf bamboo competition, while rodent predation is heavier beneath a cover
of bamboo (Wada, 1993; Abe et al., 2001). Even so, severe browsing may destroy regeneration not protected in fenced enclosures
(Akashi, 1997), or unless the deer population is reduced (Akashi and
Nakashizuka, 1999). Countermeasures might include controlled
hunting or fencing, coupled with site preparation and reproduction method cutting across large areas at one time (Sage et al., 2003;
Baumhauer et al., 2005).
2.3. Key ecological characteristics of the Fagus species related to
regeneration measures
F.s., F.c., F.g., and F.o. have shade-tolerant, dominant climax traits
that must be considered when planning regeneration projects.
Those similarities are central to the “model genus” concept in Fagus.
First, several years may pass between masting events, and these
cannot be forecast. Second, pollen effectively disperses only about
100 m, or less in closed stands. Third, seeds fall within 20 m of a
parent tree. Birds or mammals may carry the nuts farther, but that
dispersal is not dependable. Fourth, beech seedlings are sensitive
to frost, drought, and animal predation and may suffer from herbaceous competition. Fifth, beech is very shade tolerant and develops
best into high quality saplings at sites with dense regeneration and
a moderate canopy shelter, at least until the seedlings are large
enough for release. Yet those conditions reduce chances for regenerating more light-demanding associated species and increasing
tree species diversity.
3. Silvicultural systems and trends in silviculture
Effective silviculture accounts for landowner needs (Nyland,
2002) and prescribes treatments that address specific management
goals. As Dengler (1930) stated, silvicultural systems must also take
cognizance of ecologic and bio-physical features of a site and the
species of interest, i.e. key ecological characteristics.
3.1. Aims in beech management
Except for F.g., beech species are commonly the dominant component of a stand. This often encouraged managers to maintain F.c.
2177
and F.s. in pure stands, using coppice systems for fuelwood production (Kamitani, 1986; Mormiche, 1981 cited in Peters, 1997)
and high-forest systems for growing high quality sawtimber. The
alternatives of converting beech forests into plantations of fastergrowing conifers (e.g., Cryptomeria and Chamaecyparis in Japan;
Picea and Pinus in Europe) often increased snow and wind breakage,
and bark beetle infestations.
Today’s objectives include sustaining multiple services and
values from beech forests (Fujimori, 2001; Haynes et al., 2003),
often by mimicking the natural dynamics of unmanaged stands,
increasing native tree species and their mixtures, and diversifying structures between and within stands (Hahn et al., 2005;
Mohadjer, 2005; Wagner and Lundqvist, 2005; Puettmann and
Ammer, 2007). Consistent with this, several European countries
have major afforestation programmes using beech (e.g., Madsen
et al., 2005; Weber, 2005); especially for replacing off-site conifer
plantations to restore a more “natural” condition (Spiecker et al.,
2004; Bradshaw, 2005; Stanturf, 2005; Hansen and Spiecker, 2005).
Further, many researchers propose emulating gap disturbance of
unmanaged temperate forests, and that fits the regeneration characteristics of Fagus.
Within northeaster North America, beech bark disease makes
long-term management of F.g. impractical (Nyland et al., 2006b).
Landowners can maintain F.g. in stands free of beech bark disease or with apparently resistant or tolerant trees (Bogenschütz,
1983), using light partial cutting to promote small beech into
overstory positions (Ostrofsky and Houston, 1988). Yet research
has not provided effective means for identifying truly resistant
clones (Houston, 2003; McCullough et al., 2000), and that complicates management in yet unaffected areas. Among infected stands,
understory beech must be controlled in order to successfully regenerate other species (e.g. sugar maple) using shelterwood method
and selection system cuttings (Kelty and Nyland, 1981; Ray et al.,
1999; Nyland et al., 2006b; Nyland, 2008).
3.2. Regeneration methods
3.2.1. Natural versus artificial regeneration of beech
Managers commonly rely on natural regeneration in beechdominated forests having an adequate seed source, but may also
use direct seeding or planting to introduce new species and
provenances. While natural regeneration is considered the least
expensive means (Wagner and Lundqvist, 2005), it may fail. In addition, overstory trees left at low density during a long regeneration
period may decline in quality and value (Hahn et al., 2005).
Artificial regeneration is appropriate in stands lacking seed trees
of desired species or having ones not adapted to a site. In Europe,
this is relevant to afforestation programmes on former agricultural
land, for advance planting of beech beneath conifer stands, and
when a change in beech provenance is required.
3.2.2. Artificial regeneration methods
Artificial regeneration with beech is mostly by planting (e.g.,
Spiecker et al., 2004; Wagner and Lundqvist, 2005) using bare-root
undercut seedlings transplanted from nurseries at 1 to 4 years of
age. In Germany, use of wildings has gained popularity (Wagner
and Lundqvist, 2005). Container stock seedlings also offer an alternative, including use of 3 to 6 month old stock (Madsen et al.,
2006). With beech and oak, container seedlings are more resistant
to handling damage and can be used to extend the planting season
(Kerr, 1994). Direct seeding is less expensive than these planting
methods, and can be used to establish densely stocked regeneration with natural taproots (Bullard et al., 1992; Baumhauer et al.,
2005). These grow similar to planted nursery stock (Ammer and
Mosandl, 2007). Yet direct seeding requires careful handling of the
beechnuts, appropriate seedbed preparation, and careful site selec-
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tion (Ammer et al., 2002; Löf et al., 2004; Madsen and Löf, 2005).
Baumhauer et al. (2005) and Madsen et al. (2006) also recommend
using species mixtures and applying the sowing to large areas.
3.2.3. Natural regeneration methods
Taking all ecological characteristics of the genus into account,
to succeed with Fagus, natural regeneration methods must
leave a dense shelter of old trees and require a long regeneration period. With F.g., light partial cutting of single trees
or small groups has resulted in an undesirable domination
by the beech (Nyland et al., 2006b), i.e. it was very successful for beech regeneration. Similarly, for F.s., those treatments
will not ensure a diversity of other species, leading to management problems where the associated ones have important
commodity and environmental protection values (Collet et al.,
2008).
Shelterwood method with short regeneration periods should
ensure a higher species diversity (Kelty and Nyland, 1981; Ray
et al., 1999) and result in a cohort density appropriate to
high value timber production. However, with F.o., wet autumns,
mild winters, cold spring weather, late frosts, frequent droughts,
and an acidic humus make success with shelterwood method
uncertain (Sagheb-Talebi et al., 2005). With F.s., difficult abiotic
conditions, seed-eating animals, and interfering ground vegetation have sometimes caused failures. With F.c., shelterwood
method will fail where dwarf bamboo and shrub species interfere with the tree regeneration. Generally, regenerating F.g. is not
a goal in the aftermath forests of North America (Nyland et al.,
2006b).
To this end, there virtually is no single cutting treatment that
simultaneously fulfils recent management aims like diversity of
species, high quality of young beech, and naturalness with regard
to beech dominated natural ecosystems.
4. Some opportunities for regeneration research
No single silvicultural system addresses all ecologic and managerial factors that influence modern beech forestry. Site conditions
vary among stands, and different kinds of forest services often
depend on maintaining a variety of structural conditions in neighbouring parts of a forest and its adjacent landscape. That may
necessitate use of more than one silvicultural system within
a forest and across ownerships. To that end, research should
continue to explore the responses that follow a wide range of
stand treatments, to further articulate the possibilities available to
landowners.
Understory root suckers of F.g. have reduced shrubs, herbs,
and tree regeneration in areas infected with beech bark disease (Forrester and Bohn, 2007), resulting in a gradual increase
of beech in aftermath stands. The developing F.g. trees die at
pole stage, and are replaced by new root suckers, further promoting beech dominance (Nyland et al., 2006a). Research should
investigate the resulting stand development pathways and the
implications of reduced bio-diversity and simplified structural
complexity in affected stands. In addition, field trials should continue to explore non-herbicide methods for controlling F.g. to
insure favourable species diversity in new cohorts. Managers also
need field-expedient methods for identifying tolerant and resistant
beech clones to manage for plant species diversity and continued
mast production.
Young seedling-origin beech grows more slowly than many
associated species (Beaudet and Messier, 1998; Ray et al., 1999;
Dreyer et al., 2005), but faster than some. Growth curves for F.s.
allow managers to compare their development at intermediate
ages, but not during early stages of cohort development when
cleaning might seem appropriate, as with oaks that need early
release to keep them free of oppression by beech (von Lüpke, 1998).
This matter requires further attention in research, particularly in
regions where global change might alter environmental conditions
in ways that affect the relative growth of different species, and
where continued beech dominance seems unwise.
Better understanding of the fundamental performance characteristics of trees during the regeneration phase will be needed to
insure that silviculture accommodates these potential future conditions. With F.g. and F.s. the seeds, germinants, and seedlings (Geßler
et al., 2007) are drought sensitive, potentially affecting beech
seedling regeneration at sites that might be tempered by important
climate change. In this regard, Jump et al. (2007) identified temperature as an ecological factor critical to regeneration success at the
southernmost distribution of F.s., where it seems limited by moisture stress (Aranda et al., 2000). Similarly, Löf et al. (2005) stressed
that drought strongly influences beech performance when light is
not limiting, and Bolte et al. (2007) identified a shortage in nitrogen supply among additional drought-induced stresses to beech
regeneration. In addition, Lendzion and Leuschner (2008) showed
that atmospheric vapour pressure deficits may become limiting to
beech regeneration, even with moisture in the rooting medium
near optimum. Further, with disturbance-adapted species, irradiance and temperature affect the competitive interference to beech
(Fotelli et al., 2004). Thus, gaining greater understanding of the ecophysiological traits of the several beech species and their associates
through laboratory research with seedlings has become increasingly important. Regrettably, those investigations with Fagus have
involved only F.s. to date.
Well-documented and widely used beech regeneration strategies of the past may have little value if future weather phenomena
differ importantly from those of today’s climate. To date, silvicultural research into adapting cutting regimes to accommodate
climate change has primarily looked at effects on soil moisture
(Czajkowski et al., 2005), including those caused by overstory interception, coupled with soil moisture reduction due to transpiration
from the older trees (Wagner et al., 2009). Yet ways to compensate for higher temperatures, reduced precipitation, and vapour
pressure deficits remain untested. Research must explore these
matters (Lendzion and Leuschner, 2008). Possibilities may include
regenerating F.s. in mixtures with more drought-tolerant species,
and using direct seeding or planting to introduce drought-resistant
beech provenances. However, this latter measure may increase
risks of introgression to a less well-adapted population, similar to
the genetic pollution in wild fruit tree species (for Malus sylvestris
see Wagner, 2005) and with Populus nigra (Heinze, 1998) in Europe.
These risks must be evaluated before provenance mixing becomes
common with forest tree species like beech.
Despite the historic and continuing northward expansion of
F.c. on Hokkaido Island (Peters, 1997), ecologists cannot explain
the extensive post-glacial dispersal of F.g. (Clark et al., 1998), or
how some northern outliers of F.s. became established in Sweden
(Björkman, 1999). Understanding how this dispersion occurred in
the past may help forest ecologists to forecast how climate change
might alter the future distribution of heavy seeded species like
Fagus.
Research must also assess new silvicultural options for integrating a broader array of commodity and non-market objectives, and
scrutinize recently proposed “near-to-nature” approaches. Studies
should differentiate between the nature of a silvicultural system
for influencing long-term stand development, and the component
treatments that managers might use in at a single point in stand
development. Research should also explore ways to coordinate sustained management of single stands with needs and opportunities
across an entire forest ownership, at a landscape scale, and across
ecological spans of time (Nyland, 2002).
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S. Wagner et al. / Forest Ecology and Management 259 (2010) 2172–2182
5. Conclusions
This review introduces the idea of a “model-genus” by comparing ecological characteristics of the most prominent Fagus species.
Findings validate this idea. It enabled us to (i) delineate regeneration strategies that seem appropriate to any of the species of
concern, based upon autecological characteristics; (ii) stress the
importance of management objectives in silvicultural planning;
and (iii) articulate the synecological features critical to successful
regeneration. The latter has sometimes been underestimated in the
past.
Knowledge derived from this review indicates that management
objectives have become increasingly diverse, necessitating a broadening of the silvicultural systems used in these forests. No universal
“best-practice” can be applied worldwide, even within the ecological similar genus Fagus. This demands continued research into
mixing species for high quality timber production, addressing the
decline of Fagus grandifolia in aftermath forest management, and
evaluating Fagus crenata management in coexistence with dwarf
bamboo. Global climate change presents new challenges, as the
long-distance dispersal of heavy seeded Fagus species is poorly
understood, and the drought sensitive seedlings may succumb to
extreme weather.
A model-genus approach should broaden silvicultural knowledge as scientists explore new pathways for managing forests. It
may also have relevance with other genera like Betula for illustrating the similarities and differences of species with low to
intermediate shade tolerance (Perala and Alm, 1990a,b).
Acknowledgements
The authors wish to thank the two special issue guest editors
Kazuhiko Terazawa and Koen Kramer as well as two anonymous
reviewers for their constructive comments on earlier drafts of this
manuscript. Special thanks go to Peter Meyer and Vath Tabaku
for leaving gap frequency data of Albanian beech forests to our
intended purpose.
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VICTORIA A N A T O L Y I V N A TSYGANKOVA
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University of Illinois at Chicago
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