ORIGINAL PAPER

The sprouting ability of the main tree species in Central Europeancoppices: implications for coppice restoration

Radim Matula • Martin Svatek • Jana Kurova •

Lubos Uradnıcek • Jan Kadavy • Michal Kneifl

Received: 6 September 2011 / Revised: 16 January 2012 / Accepted: 17 February 2012

� Springer-Verlag 2012

Abstract Coppicing was widespread across Europe for

many centuries, but during the last 150 years, it has been

largely abandoned. Most of the former coppices have been

converted to high forest, especially in Central and north-

western Europe. Recently, there has been renewed interest

in restoring coppices in some regions, primarily for bio-

mass production and nature conservation. However, there

is limited information on the sprouting ability of European

tree species, which is the key prerequisite for successful

coppice restoration. To address this gap, we evaluated the

post-harvest stump sprouting of the three main species of

Central European coppices—sessile oak (Quercus petraea

(Mattuschka) Liebl.), European hornbeam (Carpinus bet-

ulus L.) and small-leaved lime (Tilia cordata Mill.)—in

relation to the stump diameter and density of residual trees.

Lime and hornbeam resprouted from stumps of all diam-

eters, but sprouting ability declined with increasing stump

diameter in sessile oak. Lime produced greater numbers of

sprouts with greater diameters and heights than either oak

or hornbeam. The number of sprouts per stump increased

with stump diameter in all three species as did the height of

lime and hornbeam sprouts, whereas there was no such

effect on the height of oak sprouts. The sprouting of

hornbeam and oak increased and decreased, respectively,

with an increasing density of residual trees. In conclusion,

our study shows that all of the studied species are able to

resprout even at an old age and after a long period of

neglect; however, there were important differences among

the species. The results also indicate that the age of the

parent trees at the time of cutting may significantly affect

the tree species composition of a newly restored coppice.

Keywords Sprouting � Coppice restoration � Coppice

with standards � Tilia cordata � Quercus petraea � Carpinus

betulus

Introduction

Coppicing, the oldest silvicultural system known in most of

the countries in the world (Matthews 1991; Fujimori 2001),

was widespread throughout Europe (Fujimori 2001;

Rackham 2003; Honnay et al. 2004; Szabo 2005) until the

second half of the 19th century (Buckley 1992; Peterken

1993). Since then, active coppice management has been

largely abandoned in response to a declining market for

coppice products (Peterken 1993; Buckley 1992; Jansen

and Kuiper 2004), and most of the former coppices have

been converted to high forest (Matthews 1991), especially

in Central and northwestern Europe (Hedl et al. 2010).

There is now an increasing debate over coppicing for

biomass production (Hall 2002; Scholz and Ellerbrock

2002; Jansen and Kuiper 2004; Nestorovski et al. 2009) or

for other purposes, such as urban forestry (Rydberg 2000;

Nielsen and Møller 2008). Much of the recent interest in

coppice restoration also arises from nature conservation

reasons and has been particularly stimulated by entomo-

logical studies that have linked a decline in insect species

Communicated by C. Ammer.

R. Matula (&) � M. Svatek � J. Kurova � L. Uradnıcek

Department of Forest Botany, Dendrology and

Geobiocoenology, Faculty of Forestry

and Wood Technology, Mendel University in Brno,

Zemedelska 3, 613 00 Brno, Czech Republic

e-mail: radim.matula@mendelu.cz

J. Kadavy � M. Kneifl

Department of Forest Management, Faculty of Forestry

and Wood Technology, Mendel University in Brno,

Zemedelska 3, 613 00 Brno, Czech Republic

123

Eur J Forest Res

DOI 10.1007/s10342-012-0618-5

diversity to the abandonment of coppicing (e.g. Warren

1987; Benes et al. 2006; Freese et al. 2006; Spitzer et al.

2008). Geographically, most of the efforts to restore cop-

pices have been made in Britain (Peterken 1993; Rackham

2003; Joys et al. 2004), but re-coppicing is currently being

considered in several Central, southeastern (Vacik et al.

2009; Wolfslehner et al. 2009) and Southern (Coppini and

Hermanin 2007) European countries.

Because coppicing involves cutting trees near ground

level and allowing them to regrow, the key factor in suc-

cessful coppice restoration, whether for nature conserva-

tion or for economic reasons, is the ability of trees to sprout

from the cut stump. In general, woody plant species are

divided into two groups: sprouters and non-sprouters (Bond

and Midgley 2001; Vesk 2006). However, there is a large

group of species that behave as non-sprouters under

favourable site conditions with no or little disturbance, but

in poor or severely and frequently disturbed sites may use

sprouting as their dominant regeneration mechanism

(Bellingham and Sparrow 2000; Bond and Midgley 2001;

Del Tredici 2001). Although variation in the sprouting of

tree species has been reported from several vegetation

types and disturbance regimes around the world (Lamson

1988; Kays and Canham 1991; Del Tredici 2001; Atwood

et al. 2009), there is still very little information in the

literature on sprouting ability in the main European tree

species (especially in relation to tree size/age). The only

studies available are on birch (Betula pendula L.), willow

(Salix spp.) and European aspen (Populus tremula L.) in

Northern Europe (Hytonen 1994; Rydberg 2000; Johansson

2008; Hamberg et al. 2011); Holm oak (Quercus ilex L.;

Ducrey and Turrel 1992; Retana et al. 1992) and chestnut

(Castanea sativa Mill.) (Giudici and Zingg 2005) in

Western Europe; and some macchia species in Southern

Europe (Giovannini et al. 1992). To the best of our

knowledge, there has not been a study investigating the

sprouting ability of the main tree species in Central Euro-

pean coppices, such as sessile oak (Quercus petraea

(Mattuschka) Liebl.), hornbeam (Carpinus betulus L.) or

lime (Tilia spp.).

In Britain, very little empirical data exist to assess the

effectiveness of recent coppice restoration projects (Joys

et al. 2004), but some authors have reported problems with

sprouting from older stools that had not been cut for dec-

ades (Fuller and Warren 1993; Peterken 1993). This age

dependence of sprouting may be crucial for the successful

restoration of coppices because the potential sites for res-

toration are either high forests or abandoned coppices,

neither of which has been cut for many years. Numerous

studies examining the relationship between tree age and

sprouting can be found throughout the North American

literature, in which most authors have observed a decline in

stump sprouting as the diameter and age of the parent tree

increases (Johnson 1977; MacDonald and Powell 1983;

Dey and Jensen 2002; Sands and Abrams 2009), although

they have also found important differences in sprouting

abilities among species (Kays et al. 1988; Atwood et al.

2009). Some European studies (Burley et al. 2004; Utınek

2004) claim that the ability of trees to resprout from the

stump generally decreases with age, but without any data

supporting this hypothesis. Rackham (2003) noted that

many examples refute the commonly held belief that

coppice stools will not sprout if they were last cut more

than 40 years ago.

This study investigated and compared the resprouting of

three principal tree species in Central European coppices—

sessile oak (Quercus petraea (Mattuschka) Liebl.), Euro-

pean hornbeam (Carpinus betulus L.) and small-leaved

lime (Tilia cordata Mill)—to assess the possibility of

converting the high broadleaved forests of Central Europe

to coppices and coppices with standards, two short-rotation

systems that were practised in the area in the past. Spe-

cifically, the objective of this study was to determine the

interspecific differences in (1) the probability that a tree

would produce stump sprouts after it was felled and (2) the

initial sprout growth, both in relation to the stump diame-

ter/age and to the density of the residual trees. Residual

timber trees, or standards, used to be commonly left

standing in the coppices with standards, but data on the

effect of standard density on the sprouting of coppiced

trees are missing in the European literature. Therefore, the

present study was carried out not only in plots from which

all of the trees had been harvested (i.e. coppices) but also in

coppices with standard trees. Residual trees limit the

amount of light that reaches the forest floor and may

therefore either favour or hinder the regeneration of dif-

ferent tree species (Ausden 2007). We hypothesised that by

varying the density of residual trees, it would be possible to

support or suppress the sprouting of some tree species and

thus manipulate the species composition of a newly

restored coppice with standards.

Methods

This study was carried out in the Training Forest Enterprise

Krtiny of Mendel University in Brno, located in south-

eastern Czech Republic (16�4005500E, 49�1303000N). The

elevation of the study area is 401 m a.s.l. The bedrock is

chalk, and the soils are brown forest soils with a high

calcium content. The average annual rainfall is 510 mm,

and the average annual air temperature is 8.4�C. The

average temperature in July (the warmest month) is 18.4�C,

and the average temperature in January (the coldest month)

is -2.1�C based on data from 1960 to 2010 from the Brno

weather station.

Eur J Forest Res

123

The study area was an active coppice for at least

200 years in the eighteenth and nineteenth centuries and

was documented as an active coppice as late as 1898

(Kadavy et al. 2011). However, from 1902 to 1920, the

coppice underwent a transformation to a high forest (Ka-

davy et al. 2011) and was kept as a high forest until January

2009 when 4 ha of the forest was harvested with an

intention to restore a short-rotation coppice system. Prior to

this harvest, all of the trees with diameter at breast height

(DBH) C7 cm were identified to the species level, and their

exact positions were recorded using the Field-Map tech-

nology (IFER, Ltd., Jılove u Prahy, Czech Republic; for

details of the technology see Hedl et al. 2009) so that they

could be easily located after they were cut. The studied

forest had a total basal area of 33.2 m2 ha-1 (BA) with 689

trees/ha with a DBH C 7 cm and was dominated by sessile

oak (Quercus petraea (Matt.) Liebl.), small-leaved lime

(Tilia cordata Mill.) and European hornbeam (Carpinus

betulus L.). The sprouting of these three species was

studied in eight experimental square plots of 2500 m2 each

that were randomly placed within the 4 ha restored cop-

pice. In all of the plots, the trees were cut approximately

5–10 cm above ground level. The season of harvest and the

density of residual standing trees followed the basic man-

agement practices that were common in the region

120 years ago based on the historical management plans of

the Training Forest Enterprise. To study the effect of the

density of residual trees on sprouting and sprout growth,

four densities of healthy canopy trees of sessile oak were

left uncut, each density in two plots. The four densities

used were 0 (i.e. clear-cut), 20 (1.1 m2 in BA), 35 (1.8 m2

BA) and 50 (2.5 m2 BA) trees per plot. The residual trees

averaged 21.1 m in height and 41.5 cm in DBH. The whole

4 ha stand was fenced because there was significant game

pressure in the area. The fence was checked at least once

every other week because the game animals had damaged it

several times.

A year after the cutting, in winter 2009/2010, we

checked every stump of the studied species in all of the

plots to determine whether the stump had produced at least

one live sprout. In total, we evaluated 321 lime stumps, 315

oak stumps and 310 hornbeam stumps in the 8 plots. On the

stumps that sprouted, we counted all of the sprouts and

measured the height and diameter (at a height of 1 cm

above the sprout base) of the 5 tallest sprouts within each

stump. Although the fence around the study area was

checked regularly, some game managed to break in and

browse some of the sprouts. Therefore, all of the stumps

whose sprouts showed signs of browsing were evaluated

only in terms of whether they had sprouted or not, and the

respective sprout measurements were left out of the data

analysis. We measured the diameter of each stump in two

perpendicular directions, and the final diameter was defined

as the average of the two diameters. We also revisited the

unsprouted individuals of the first year in the middle of the

vegetation season in summer 2010 to check whether they

had resprouted in the second year. However, there were

almost no newly sprouting individuals (only 3 hornbeam

stumps), so we pooled the data from the first and the second

years into one variable (sprouted = 1, not sprouted = 0).

We divided stumps into three diameter classes and calcu-

lated tree rings on 33 randomly selected stumps of lime and

33 stumps of oak within each diameter class (i.e. 99 per

species) to test whether stump diameter could be used as a

predictor of the age of parent trees. The tree ring counting

could not be completed for hornbeam because its tree rings

were not visible enough to provide reliable age estimates.

We carried out a regression analysis to evaluate the

relationship between the age and stump diameter. To

determine whether the density of residual trees, stump

diameter or an interaction between these two variables had

an effect on tree stump sprouting, we used generalised

linear models (GLM) with a binomial error distribution

(link logit). The effects of the density of residual trees,

stump diameter and species, including their mutual inter-

actions, on the number of sprouts produced per stump were

tested using a GLM with a quasi-Poisson error distribution

(link identity). The species were coded by dummy vari-

ables by orthogonal contrasts. The quasi distribution was

used because of the presence of overdispersion. To test the

effects of the density of residual trees and stump diameter

on the number of sprouts within each species, we used the

same GLM with a quasi-Poisson error distribution.

We used general linear models (OLS) to test the effects

of stump diameter, the density of residual trees and species

on the height and diameter of the tallest sprouts. Using the

same method, we tested the effects of stump diameter and

the density of residual trees on the height and diameter of

the tallest sprouts within each species. We carried out the

data analyses using both the diameter and height of the

single tallest sprout within a stump as well as with the

mean height and diameter of the 5 tallest sprouts within a

stump. In all of the cases, the models that were constructed

using the mean of the 5 tallest sprouts showed very similar

results but with a better fit than the models using only the

tallest sprout. Thus, for simplification, we present here only

the results for the mean values here.

We performed a backward variable elimination from the

maximal model to select the final models. The final models

were chosen on the basis of the highest Radj2 (for OLS), the

lowest P values and the lowest Akaike’s information cri-

terion (AIC). The generalised R2 coefficient of determi-

nation (Nagelkerke 1991) was computed for GLMs. When

necessary, the data were log-transformed to the base 10. To

test the differences between species, and between residual

tree densities in the mean number of sprouts, sprout heights

Eur J Forest Res

123

and diameters, we used an ANOVA with a Unequal N

Honestly Significant Difference (HSD) post hoc test. All of

the analyses were performed in the R2.12.0 statistical

environment (R Development Core Team 2010). The

ggplot2 package (Wickham 2009) was utilised to visualise

the results.

Results

There was a very strong linear relationship between the

stump diameter and the age of the parent tree in both oak

(Radj2 = 0.95; F = 1276.1; P \ 0.0001; log(y) = 1.084 ?

1.162log(x)) and in lime (Radj2 = 0.87, F = 622.2, P \

0.0001; log(y) = 0.971 ? 1.397log(x)).

Sprouting probability

Of the 321 lime stumps, only 1 did not sprout, and there-

fore, this species was not considered in the analysis of

sprouting probability. On average, 93.8% (±3.21) of the

hornbeam stumps and 61.1% (±5.23) of the sessile oak

stumps produced sprouts. There was a significant decrease

in probability of sprouting with an increase in stump

diameter in sessile oak (v2 = 11.60, df = 1, P \ 0.0001;

Fig. 1a) as well as with an increasing number of residual

trees (v2 = 5.12, df = 1, P = 0.024; Fig. 1b); the inter-

action of these two variables was insignificant (v2 = 1.08,

df = 1, P = 0.296). In hornbeam, the probability of

sprouting increased with an increase in stump diameter

(v2 = 6.60, df = 1, P = 0.029; Fig. 1a) as well as with an

increase in the number of residual trees per plot (v2 = 6.34,

df = 1, P = 0.013; Fig. 1b). As in oak, the interaction of

these variables was insignificant (v2 = 0.01, df = 2,

P = 0.948).

The number of sprouts

The number of residual trees, stump diameter, species and

an interaction of species with stump diameter had a sig-

nificant effect on the number of sprouts per stump (Nage-

lkerke’s R2 = 0.48, P \ 0.0001). The most significant

Fig. 1 The probability of

resprouting after harvest in

relation to a stump diameter and

b the density of residual trees.

The line shows the predicted

relationship from the

generalised linear model using a

binomial error distribution

Eur J Forest Res

123

variable was stump diameter (F = 205.41; df = 1;

P \ 0.0001) followed by species (F = 81.03; df = 2;

P \ 0.0001), the number of residual trees (F = 11.03;

df = 2; P \ 0.001) and the two-way interaction between

stump diameter and species (F = 6.59; df = 2;

P = 0.002). The number of residual trees in an interaction

with the other variables did not affect the number of

sprouts (P [ 0.05).

The maximum numbers of sprouts produced per stump

were 411 in lime, 289 in oak and 161 in hornbeam. There

was not a significant difference in any of the species in the

mean number of sprouts per stump in the plots without

residual trees (Table 1). However, in the plots with a

density of 25 and 35 residual trees per plot, the lime

stumps had approximately twice the number of sprouts as

the hornbeam and oak stumps, and the difference was even

greater in the plots with 50 residual trees (Table 1).

In hornbeam, only stump diameter had a significantly

positive effect on the number of sprouts (Fig. 2), and the

density of residual trees was not significant either by itself

or in an interaction (Table 2). In oak, stump diameter and

the density of residual trees in an interaction with stump

diameter had an effect on the number of sprouts, whereas

the density of residual trees by itself was insignificant

(Table 2; Fig. 2). In lime, the number of sprouts increased

with an increase in stump diameter as well as with an

increased density of residual trees (Table 2; Fig. 2). The

interaction of the density of residual trees with stump

diameter was also significant (Table 2) in this species.

Sprout height

Lime had the greatest mean sprout height, and oak had the

lowest (Table 1). Species, stump diameter and the density

of residual trees had a significant effect on the mean

maximum sprout height (Radj2 = 0.32; df = 2; F = 39.63;

P \ 0.0001). Species was the most significant factor

(F = 97.05; df = 2; P \ 0.0001) followed by the effects

of stump diameter (F = 87.63; df = 1; P \ 0.0001) and

the density of residual trees (F = 4.53; df = 1; P \0.0001). All of the interactions were not significant

(P [ 0.05).

In hornbeam, the mean maximum sprout height signif-

icantly increased with stump diameter and decreased with

an increase in the density of residual trees (Fig. 3), but the

interaction between these two variables was insignificant

(Table 2). No significant effects of either stump diameter

or the density of residual trees on the maximum shoot

heights were found in oak (Table 2). In lime, the mean

maximum height significantly increased with increases in

the stump diameter and the density of residual trees

(Fig. 3), whereas the interaction between these two vari-

ables was insignificant (Table 2). Ta

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Eur J Forest Res

123

Sprout diameter

Lime had a significantly greater mean maximum sprout

diameter than both oak and hornbeam (Table 1). Species,

stump diameter and the density of residual trees proved to

have significant effects on the sprout diameter of the 5

tallest sprouts (Radj2 = 0.48, F = 98.61, P \ 0.0001). None

of the interactions were significant (P [ 0.05). The species

variable had the strongest effect (F = 275.49, df = 2,

P \ 0.0001); the effects of stump diameter (F = 31.80,

df = 1, P \ 0.0001) and the density of residual trees

(F = 5.19, df = 1, P = 0.023) were much weaker.

In hornbeam, only the density of residual trees had a

significant effect on the sprout diameter of the tallest

sprouts (Table 2; Fig. 4). In oak, no explanatory variable

affected the sprout diameter of the tallest sprouts (Table 2;

Fig. 4). In lime, only the stump diameter had a significant

effect on the sprout diameter (Table 2; Fig. 4).

Discussion

As expected, we found that stump diameter is a reliable

linear predictor of the age of the parent tree. Because this

stump diameter/age relationship was highly significant in

both lime and oak, we presume that a similar relationship is

likely to exist in hornbeam as well.

Our study has shown that it is possible to convert a

Central European high broadleaved forest into a coppice or

coppice with standards because all three species studied

proved to have a good ability to resprout from the wide

range of observed stump diameters. However, there were

clear differences in the sprouting abilities of the three

species. In previous studies (Del Tredici 2001; Vesk and

Westoby 2004), the ability to resprout after cutting has

been generally considered to decline with increasing age

and stump diameter in most tree species. From a practical

perspective, the ability of stumps to sprout may be the main

obstacle for coppice restoration in Central Europe because

many of the potential candidates for restoration are either

abandoned or transformed coppices that were last cut

several decades ago. Nevertheless, our results showed that

the probability of sprouting increased or remained consis-

tently high with an increased parent tree age and diameter

in lime and European hornbeam. Only sessile oak had a

significant nonlinear decrease in sprouting probability from

approximately 80% for the smallest and youngest trees to

less than 40% for the trees with the largest stumps, which

were approximately 90 years old. Weigel and Peng (2002)

showed a similar decline in the probabilities of sprouting

with increasing parent tree age in five North American oak

species.

In our study, the relatively low sprouting probability of

the older stumps of sessile oak contrasts with an almost

100% sprouting potential of lime across all of the ages and

diameters of the parent trees. European hornbeam proved

to have a very high sprouting ability but with a different

pattern from the other two species. The hornbeam stumps

demonstrated a steady increase in their sprouting ability

from lower values in the youngest and smallest trees to

almost 100% sprouting of the stumps of the oldest and

biggest trees. The difference in sprouting among the tested

species may be related to differences in bark thickness, as

was shown by Wilson (1968), who linked a failure of

hidden epicormic buds to develop into new sprouts to the

physical resistance of the bark. As stump diameter

increases with age, the bark thickness increases and so does

its physical resistance, which may lead to the higher

sprouting failure rates of older trees (Johnson et al. 2002).

This dependence upon bark characteristics may be one of

the causes of the comparably better sprouting of old lime

and hornbeam trees, which both maintain a thin and soft

bark throughout their life spans. In contrast, the bark of

sessile oak becomes significantly thicker and harder to

penetrate with an increase in stem diameter. The other

possible reason for the interspecific differences in sprouting

could be the resource economy of tree species (Clarke et al.

2010). Some sprouters may allocate more resources to root

than shoot biomass and store large non-structural carbo-

hydrate reserves to support bud growth (Clarke et al. 2010),

Fig. 2 The relationship

between the number of sprouts

per stump and stump diameter.

The line shows the prediction

based on generalised linear

models with a Poisson

distribution. The pointsrepresent individual

observations

Eur J Forest Res

123

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Eur J Forest Res

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but this fact has been observed mainly in herbaceous and

shrub species while data on tree species are largely lacking

(Bond and Midgley 2001; Clarke et al. 2010). The present

study has shown that lime produced more sprouts than oak

and hornbeam. In addition, the lime sprouts were taller and

thicker than those produced by oak and hornbeam, indi-

cating that lime produced more biomass than the other two

species in the first year. These facts coupled with an almost

100% sprouting probability of lime stumps demonstrate

that lime is the best sprouter of the studied species. A very

high sprouting ability of small-leaved lime has been pre-

viously reported by several authors (Pigott 1989; Rackham

2003); however, no empirical data on this topic had been

published. The vigorous sprouting of lime is even more

interesting when its ability to regenerate by seeding is also

taken into account. In general, the regeneration strategy of

woody plant species is considered to be a trade-off between

seeding and sprouting (Bellingham and Sparrow 2000;

Bond and Midgley 2001, 2003). Nonetheless, our study

showed that some species, such as lime, may possess both

well-developed regeneration modes. In addition to being a

very good sprouter under a severe disturbance regime, a

mature lime tree produces thousands of viable seeds every

year when allowed to reach maturity. Our study area might

be one example of such a regime because it contained

abundant lime seedlings and saplings (Matula and Urad-

nicek, unpublished data) that originated from the seeds of

the same parent trees whose stumps vigorously sprouted

after they were cut. The absence of a trade-off between

sprouting and seeding was most apparent in the case of

lime, but we also did not find any evidence of such trade-

offs in oak or in hornbeam. In addition to the sprouting

stumps, there were thousands of seedlings of the two spe-

cies in the study plots (Matula and Uradnicek, unpublished

data).

Young stump sprouts benefit from the mature root sys-

tem of a parent tree, which allows them to quickly out-

compete and outgrow seedlings and seedling sprouts

(Johnson et al. 2002). It is evident that post-harvest

sprouting rate of nearly 100% of lime and hornbeam

stumps, compared with only 61% resprouting of oak

stumps, will result in a shift in tree species composition in

favour of lime and hornbeam even though the stump

sprouts themselves may not occupy all of the growing

space (Sander et al. 1984). Our results also indicate that an

older age of the stand at the time of cutting will result in a

more pronounced shift towards species other than oaks.

However, oak is one of the most common species in

today’s old abandoned coppices in Central Europe, indi-

cating that there was a mechanism that allowed coppiced

oak to outcompete other tree species. A short-rotation

period was probably one of the most important mecha-

nisms. Coppicing was practised in the region typically on a

5–20-year rotation (Cotta 1856; Polansky 1947), which is

Fig. 3 The relationship

between the mean height of the

5 tallest sprouts per stump and

stump diameter (cm). The lineshows the prediction based on a

regression analysis, and the

points represent individual

observations

Fig. 4 The relationship

between the mean diameter of

the 5 tallest sprouts per stump

and stump diameter (cm). The

line shows the prediction based

on a regression analysis, and the

points represent individual

observations

Eur J Forest Res

123

approximately the age of the smallest stumps of sessile oak

in our study that, according to our findings, have a high

probability of sprouting. As for lime, Rackham (2003)

suggested that on a coppice rotation shorter than 15 years,

a lime tree has very little chance to flower and produce

seeds, and this limitation may have prevented lime from

expanding in British coppices. The other possible factor

may have been that lime produces a comparatively lower

quality wood than oak. Historical documents on forest

management in the study area, some of which date back to

the 16th century, usually mention oak and less often

hornbeam as the species present in coppices, but there is

not a single word about lime. We think that lime may have

been considered to be an undesirable species and therefore

removed from coppices to support the production of oak

wood.

Our findings indicate that the effects of stump diameter

on the initial development of sprouts differ among the

species. Although in all three studied species the number of

sprouts per stump significantly increased with stump

diameter, only in lime trees did stump diameter represent

the major source of variation in the number of sprouts,

sprout height and sprout diameter. This finding suggests

that bigger and therefore older lime parent trees produce

taller and larger diameter sprouts in larger quantities than

younger parent trees after they are cut down, but in oak and

hornbeam, there are other more important factors, which

were not captured in this study, that influence their initial

sprout growth. These findings are inconsistent with the

results of most North American studies on oak sprouting,

which have shown a decline in the number of oak sprouts

and the mean annual growth associated with increasing

stump diameter (e.g. Johnson 1977; Weigel and Peng 2002;

Sands and Abrams 2009). However, our findings support

the observations of Kays et al. (1988) and Atwood et al.

(2009), who found significant differences in sprouting

abilities among oak species.

There were also clear differences in the effect of the

density of residual trees on the stump sprouting among the

species, although this effect was much weaker than that of

stump diameter. Our results indicate that in hornbeam, the

chance that a tree will resprout after harvest increases with

an increasing density of residual trees, whereas there is an

opposite effect in oak. Lime stumps produced a greater

amount of sprouts and taller sprouts under higher densities

of residual trees, whereas in hornbeam, the height and the

diameter of sprouts decreased with an increasing density of

residual trees. Thus, although the parent tree diameter and

age itself have greater effects on the sprouting of the

studied species, foresters may to some extent influence the

sprouting probability and initial sprout growth of oak,

hornbeam and lime in a newly restored coppice with

standards by varying the density of residual trees.

Lime and hornbeam are strong competitors to oak in

nutrient-rich soils but are rare in nutrient-poor sites where

oak usually dominates. However, the significant shift from

nutrient-poor to rich mesic sites in the past century that

has been documented in the lowland forest of Central

Europe (Hedl et al. 2010) may make many previously

unsuitable sites favourable for both lime and hornbeam. In

addition, the easily dispersed seeds and shade tolerance of

lime and hornbeam allow them to establish high levels of

advanced regeneration even in stands without any mature

individuals of these species. In contrast, oak is a light-

demanding species even at early regeneration stages, so a

closed canopy usually does not allow its advanced

regeneration to grow or even survive. Therefore, when the

stand is harvested, the ability of lime and hornbeam to

regenerate vigorously by sprouts, seedlings and previously

established saplings gives both species a competitive

advantage over the co-occurring oak. These differences in

regeneration dynamics must be considered when planning

a coppice restoration because many of the potential stands

for such restoration in Central Europe are dominated or

co-dominated by sessile oak. The re-coppicing of those

stands may change their composition in favour of better-

sprouting and shade-tolerant species such as lime and

hornbeam.

Conclusions

Our study has important practical implications for coppice

restoration. We showed that all of the main tree species in

Central European coppices are able to resprout even at an

old age and after a long period of neglect; however, there

were important differences among species. Further

research is needed to determine which factors (such as

resource allocation) might be responsible for interspecific

differences in resprouting, but the commonly held view

that the sprouting ability of trees generally declines with

age needs to be re-evaluated. The results also demonstrated

that both the stump diameter and the age of parent trees at

the time of cutting may significantly affect the tree species

composition of a newly restored coppice, which is of great

importance for both conservation- and economic-driven

coppice restoration projects.

Acknowledgments We thank Michal Kuchta and Martin Juhn for

their collaboration on this study. Also, we are thankful to two

anonymous reviewers for valuable comments on a previous version of

this manuscript. This study was funded by a research grant from the

Ministry of the Environment of the Czech Republic SP/2d4/59/07 for

the project ‘‘Biodiversity and target management of endangered and

protected organisms in coppices and coppice-with-standards under

system of Natura 2000’’ (TARMAG 2000), by the project NAZV CR

No. QH71161: ‘‘Coppice and Coppice-with-standards—Adequate

Forest Management Alternative for Small and Middle Forest

Eur J Forest Res

123

Owners’’, by an IGA project of the Faculty of Forestry and Wood

Technology of Mendel University in Brno titled ‘‘Use of genetic

information in forest botany, tree physiology, dendrology and geo-

biocoenology’’ and by the Institutional Research Plan MSM

6215648902/04/01/01 of the Faculty of Forestry and Wood Tech-

nology MENDELU Brno.

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