www.elsevier.com/locate/foreco

Forest Ecology and Management 237 (2006) 353–365

Intensive game keeping, coppicing and butterflies:

The story of Milovicky Wood, Czech Republic

Jiri Benes a, Oldrich Cizek a,b, Jozef Dovala c, Martin Konvicka a,b,*a Department of Ecology and Conservation, Institute of Entomology, Czech Academy of Sciences, Branisovska 31,

370 05 Ceske Budejovice, Czech Republicb Department of Zoology, School of Biological Sciences, University of South Bohemia, Branisovska 31,

370 05 Ceske Budejovice, Czech Republicc Jilesovska 3, 747 92 Haj ve Slezsku, Chabicov, Czech Republic

Received 24 January 2006; received in revised form 24 July 2006; accepted 28 September 2006

Abstract

While transfers of formerly coppiced or grazed woodlands into shady high forests cause severe declines of woodland butterflies across Europe,

increasing numbers of wild ungulates contribute to maintaining stand openness. To disentangle the relative effects of management and ungulates,

we studied butterfly assemblages in the Milovicky Wood, southeastern Czech Republic. After centuries of short-rotation coppicing, the wood was

abandoned in the 1950s and two game parks, for deer and mouflon, were established there in the 1960s. Comparisons of historical and recent

records show severe declines, but the wood still hosts 83 butterfly and burnet species, including 19 nationally endangered ones. Recording along

fixed transects disentangled effects of game keeping and management. Stands situated in the mouflon park hosted fewer species than those in either

the deer park or outside of the parks. Clearings, coppice, coppice with standards and rides hosted more species than closed forest. The strongest

predictors of composition of butterfly assemblages were plant communities and stand management, followed by vegetation covers, plant species

richness and kind of game (mouflon, deer, none). Both game and management exhibited independent effects. Past high game densities contributed

to butterfly losses, but have maintained open structures absent from woods managed for timber. Under reduced densities, mouflon exhibit adverse

effects on butterflies but deer do not. Recent plans to transfer the area to high forest are incompatible with conserving local butterflies and incur high

costs of forest protection against the animals. In contrast, re-establishment of active coppicing for fuel wood production would be optimal for

butterflies, compatible with game keeping. Finding a balance between game and traditional forms of management offers an opportunity for

threatened biodiversity of European lowland forests.

# 2006 Elsevier B.V. All rights reserved.

Keywords: Butterfly conservation; Central Europe; Coppice management; Deer; Lepidoptera; Oak

1. Introduction

Light and sparse deciduous forests of lowland temperate

Europe host a remarkable number of threatened butterflies. Five

species – Coenonympha hero, Euphydryas maturna, Leptidea

morsei, Lopinga achine and Parnassius mnemosyne – are

protected by the Habitat Directive of the European Union (92/

43/EEC), but the number of declining woodland species is

considerably higher (Van Swaay and Warren, 1999; Benes

* Corresponding author at: Department of Ecology and Conservation, Insti-

tute of Entomology, Czech Academy of Sciences, Branisovska 31, 370 05

Ceske Budejovice, Czech Republic. Tel.: +420 38 777 5312;

fax: +420 38 531 0354.

E-mail address: konva@entu.cas.cz (M. Konvicka).

0378-1127/$ – see front matter # 2006 Elsevier B.V. All rights reserved.

doi:10.1016/j.foreco.2006.09.058

et al., 2002). Open woodlands suitable for butterflies had been

maintained for centuries by historical uses, such as coppicing

and forest pasture (Warren, 1987; Buckley, 1992; Sparks et al.,

1994; Konvicka and Kuras, 1999; Bergman, 2001; Konvicka

et al., 2005). In contrast, modern high-forest management

creates shady conditions unsuitable for open-forest species

(Warren and Key, 1991; Greatorex-Davies et al., 1993;

Wahlberg et al., 2002; Freese et al., 2006; Liegl and Dolek,

2006).

In this context, understanding the effects of high game

densities on woodland butterflies increases in urgency. The

much-debated theory of wood pasture (Vera, 2000) proposes

that prehistoric woodlands had been kept relatively open first by

wild ungulates, later on by domestic animals (Hansson, 2001;

Bradshaw et al., 2003; Bakker et al., 2004; but see Birks, 2005).

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365354

Fig. 1. Map of the study area, showing position of the Milovicky Wood in

Europe, its extent (grey) and borders of the deer and mouflon parks.

Whereas woodland grazing had nearly disappeared during the

last century (Bergman and Kindvall, 2004; Saarinen et al.,

2005), numbers of wild ungulates, particularly deer, have

increased. High deer densities impede shrub and tree

regeneration (Fuller and Gill, 2001; Homolka and Heroldova,

2003; Cote et al., 2004), potentially increasing stand openness.

Such impacts should be particularly pronounced in game

parks, where high deer densities are maintained for hunting.

Having a long tradition in Europe (Rackham, 1998), game

parks contain a considerable proportion of open spaces, such as

wide glades that facilitate hunting or coppiced panels that

supply browsing and shelter for animals. Therefore, it might be

expected that game parks act as refuges for open woodland

butterflies (Stewart, 2001). On the other hand, grazing and

browsing may suppress some larval host plants and diminish

nectar supplies (Pollard and Cooke, 1994; Pollard et al., 1998;

Joys et al., 2004). Therefore, butterflies may be affected both

positively or negatively, depending on the situation (Feber

et al., 2001; Fuller and Gill, 2001). Studying the effects requires

either costly manipulation of game densities, or comparative

studies within areas that harbour high variation of game

densities and management methods.

This study attempts to disentangle the relative effects of

game keeping and stand management on butterfly richness and

community composition of the Milovicky Wood, southeastern

Czech Republic. The wood harbours a rich butterfly fauna and a

high variety of management conditions. Only parts of its

territory are used for game keeping, allowing comparison of

impacts of high and low ungulate densities, and active

coppicing is still locally practised there, unlike in other forests

in the country. The wood has been proposed as a Site of

Community Interest under the EU Habitat Directive and we

also discuss a management regime that would preserve local

butterfly fauna while being politically and economically

plausible.

2. Material and methods

2.1. Study area—history

The Milovicky Wood (Fig. 1; 488490N, 168420E, alt. 250 m)

represents the largest complex of Pannonian thermophilous

woods in the Czech Republic. It is situated within a region of

warm and relatively continental climate, at a crossroad between

the Hercynian highlands, the Carpathians and the lowlands of

Pannonia. It covers 20 km2 of rolling hills made up of limestone

and flysch and covered by quaternary deposits. Most of its

vicinity is intensive farmland, except for calcareous Palava

Hills in the West.

Humans settled the area in the Paleolithic: a classical site of

the Gravettian culture (25,000 BP, see Absolon, 1949) is just

3 km to the NW. The wood was likely coppiced since the arrival

of early farmers and the coppice cycle was extremely short for

most of recorded history, resulting in a scrubby appearance

(Vybiral, 2003). Shortly after World War II, the coppicing was

abandoned and management by singling was adopted to

improve the timber supply. Because it soon became obvious

that there was little hope for growing good timber, two game

parks covering ca. 85% of the wood were established in the

1960s: one for red and fallow deer (herein deer park: 1160 ha)

and one for fallow deer and mouflon (mouflon park: 558 ha)

(Fig. 1). The stocking peaked in late 1980s, totalling 370 deer,

380 fallow deer and 200 mouflon, with a maximum of 550

animals (1.02 animals per ha) in the smaller mouflon park.

Resulting depletion of herb layer (Chytry and Danihelka, 1993)

raised concerns among conservationists, which led to a

reduction of game densities in the mid-1990s. The current

stock is 270 animals (0.48 ha�1) in the mouflon park and 390

animals (0.34 ha�1) in the deer park.

The wood is located within the butterfly-richest grid cell of

the Czech Republic (132 species, 82% of the country’s fauna)

(Skala, 1912; Benes et al., 2002). Historically, local rarities

included Neptis sappho, E. maturna, L. morsei and L. achine.

Intensive recording had been interrupted by the establishment

of the game parks and resumed after the 1990.

2.2. Study area—current condition

A majority of the wood consists of mature (>80 years)

singled oak coppice, classified as CLOSED FOREST below.

Prevailing trees are oaks (Quercus petrea, Q. pubescens and Q.

cerris) and hornbeam (Carpinus betulus), accompanied by ash

(Fraxinus angustifoia, F. excelsior), lime (Tilia platyphylos)

and elm (Ulmus laevis). Some 140 ha (in widely scattered

panels) are coppiced as game shelters (COPPICE). Coppice

species include Acer campestre, Cornus mas, Corylus avellana,

Crataegus sp., and Sorbus torminalis.

To promote germination of oak, 200 ha of fenced

‘‘regeneration blocks’’, have been established in the last few

years. Proceeding by strips 30–50 m wide, they are sequentially

thinned to canopy cover of 10–30%. It is expected that in mast

years, oak should germinate from seeds in the thinned stands.

The thinning usually causes vigorous coppice regrowth, which

is not viewed as desirable and sometimes is suppressed using

herbicides. If not suppressed, the thinned panels structurally

resemble coppice with standards, and we regard them as such in

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365 355

this paper (STANDARDS). Following mast years, the standards

are removed in order to release generative cohort of oak. If the

seedlings fail to establish, all vegetative growth is cleared,

stumps are mechanically removed and thus prepared sites

(CLEARINGS) are planted by nursery oak. This operation may

be repeated if the nursery material fails to establish. A further

feature of the wood are wide (ca. 15 m) open rides along roads

and lanes (RIDES), where grazing suppresses woody regrowth,

maintaining low grasslands alternating with coarse grasses and

ruderal herbs.

2.3. Butterfly and vegetation sampling

Throughout this paper, ‘‘butterflies’’ refer to Papilionoi-

dea + Hesperioidea, burnet moths (Zygaenidae) and the

conspicuous diurnal moth Syntomis phegea (Arctiidae).

Nomenclature follows Karsholt and Razowski (1996). The

categorisation to woodland and steppe species follows Benes

et al. (2002) and Povolny and Gregor (1946) for butterflies and

burnets, respectively. Steppe species include all thermophilous

species, i.e. including butterflies of scrubby biotopes. Endan-

gered species follow Vrabec et al. (2006).

To compare past and present fauna (Appendix A), we used

the Czech atlas database (Benes et al., 2002) updated by

interviewing lepidopterists that recently visited the area. We

also actively searched the wood for selected priority species in

2003 and 2005. These focal surveys took 50 person-days

additional to the quantitative study described below.

For quantitative assessment, we used a modified transect

method of Pollard (1977). Because variously managed sites

were widely scattered within the wood, we did not use a regular

design with equal representation of management types. We

instead devised a 14.4 km long route zigzagging the two game

parks and stands outside of them, covering all management

types present. It consisted of 61 sections (mean length 240 m,

S.D. = 108. range 40–697 m) delimited according to stand

management. We walked the transect twice a month, from May

to August 2003 and 2004, obtaining 16 walks in total. We used a

standard pace (1.5 km/h), restricting the walks to between

9:30 a.m. and 4:30 p.m. (CE summer time) and weather

suitable for butterflies (sunny, >17 8C). If the weather

worsened, we interrupted the walk and either waited until it

improved again, or continued it next day. We alternated

direction of the walks to randomise day times of visits to

individual sections.

Butterflies were counted on per-section basis, within a 7 m

cube in front of the recorder. We used butterfly net to check

more difficult species (Benes et al., 2003a) and a semiquanti-

tative scale (<50, <100, <200, <500) to record butterflies

seen in >20 individuals. We also noted time (closest hour),

relative shade (3-point scale from full sun to shade), wind

speed (3 point scale), and nectar (3 points, none to abundant)

for each section/walk. Depending on weather, one walk took

two to four person-days.

The vegetation was surveyed twice, for spring (i.e., spring

geophytes) and early summer aspects, in mid-April and late

June, 2004. A botanist surveyed the transect for all tree, shrub

and herb species growing within a 15 m strip along it and

recorded their percentage covers, using an ordinal scale 1:

<0.01%, 2:<1%, 3:<5%, 4:<10%, 5:<25%, 6:<50% and 7:

<100%.

2.4. Variables

Length of each transect sections, time of day, nectar, shade

and wind speed are considered as covariables. The same applies

for central latitude and longitude of individual sections, plus

their quadratic terms and interaction (used to control for spatial

autocorrelation).

The explanatory variables describing game keeping and

management include: (i) game, either PRESENT (N = 33) or

ABSENT (N = 28); regeneration blocks included to the latter

category; (ii) kind of game: DEER (N = 25), MOUFLON

(N = 8) or NONE (N = 28); regeneration blocks classified as

none; (iii) management, characterising individual sections and

classified to CLEARING (N [with/without game] = 3/6),

CLOSED FOREST (N = 12/4), COPPICE (N = 4/6), RIDE

(N = 12/4) and STANDARDS (N = 2/8); (iv) surrounding

management, a continuous predictor, was obtained from

digitised aerial photographs, using ESRI ArcView GIS 3.2.

At centre of each transect section, a circle with diameter equal

to section length was drawn. Within this circle, areas subject to

individual management types were digitised, and their (arcsine

transformed) proportions of circle area were used for analyses.

The management types were as above, plus GRASSLAND and

CROP FIELD. The variables describing vegetation included:

(v) vegetation covers, distinguishing canopy (E3), shrub (E2)

and ground (E1) layers within the vegetation-sampled strips

along each transect section; (vi) spring herbs richness

(SPRING). Spring geophytes are easily recorded and may

provide a comfortable surrogate for total species richness; (vii)

higher plant species richness, split into canopy (RE3), shrub

(RE2) and ground (RE1) layers; (viii) plant community

composition, quantified by applying the principal component

analysis (PCA) for vegetation samples from individual

sections, and using the PCA scores of the sections as numeric

predictors (details in Appendix B).

2.5. Analyses

To analyse species richness, we used analysis of covariance

with section length as a covariable. Tukey’s HSD tests for

unequal N were used for subsequent comparisons. For a more

detailed analysis, we used generalised linear models (GLM,

link identity), built via stepwise addition of explanatory terms

in S-plus (MathSoft, 1999), using the Akaike information

criterion to select most parsimonious models. We standardised

all variables by subtracting their means and dividing by

standard deviations, and checked, besides of linear relation-

ships, for quadratic effects of all quantitative predictors. For

each dependent variable, we built a raw model not considering

any covariables, a covariable model based solely on selection

from covariables, and a controlled model, built by adding

explanatory terms onto the covariable model.

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365356

To study species composition, we employed two ordination

techniques in CANOCO v. 4.5 (ter Braak and Smilauer, 1998).

(i) An indirect (=unconstrained) method, the detrended

correspondence analysis (DCA), and (ii) a direct ordination,

the canonical correspondence analysis (CCA) followed by

testing the effects of environmental predictors via the Monte

Carlo permutation test.

For DCA, we used summed numbers of each species seen

per section during all walks as species data; upper values of

respective intervals were used in cases when a species was

recorded semiquantitatively. We chose a detrended analysis,

because the data exhibited an arch effect when entered as sums,

but not when entered separately for transect walks (as for CCA

below). We used detrending by segments option and log-

transformation of species counts.

Fig. 2. Numbers of all, woodland, steppe and endangered butterfly species observe

outside of game parks). The bars show mean numbers par section plus standard erro

HSD tests for unequal N).

For CCA, log-transformed species counts were entered

separately for individual walks. The temporal structure of the

data was considered in the permutation design, which permuted

transect sections (whole plots) in random and individual visits

(split plots) as cyclic shifts. We again checked for the effects of

section length and spatial position, plus nectar, time of day,

shade and wind. The latter four covariables were entered

separately per each site visit, time of day was treated in a linear,

polynomial and factorial form and the polynomial form was

used subsequently as it explained the highest amount of

variation. Next, we tested for specific effects of game keeping

(data collection: (i–ii), stand management (iii–iv) and vegeta-

tion (v–viii)). As a next step, we added game keeping (i–ii) and

management (iii–iv) to models that treated significant terms

from previous analyses as covariables. The purpose of this was

d at transect sections differing in management and kind of game (non: sections

rs. Bars with identical letters form homogeneous groups (ACOVA followed by

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365 357

to assess whether game keeping and/or management exhibited

detectable effects on butterfly composition even after fitting the

variation due to the other terms.

3. Results

3.1. Species richness

Out of 105 species (95 butterflies) reported for the wood

during the 20th century, 22 were not recorded after 1995

(Appendix A). A conservative subtraction of five species whose

occurrence is still possible (e.g. Satyrium hairstreaks) gives 17

losses, or 16% of the fauna. The losses included such

characteristic butterflies of open woodland as L. morsei, E.

maturna and L. achine. Still, we confirmed recent occurrence of

83 species, including 19 nationally threatened ones. A total of

67 species in approximately 15,000 individuals were seen along

the transect, 21 species being forest specialists and 22

representing steppe species.

Sections with game ABSENT hosted more species than

PRESENT sections (ANCOVA, F = 4.44; d.f. = 1, 58;

P < 0.05). The same applied for woodland species

(F = 11.85; d.f. = 1, 58; P < 0.01) but not for steppe species

(F = 2.57; d.f. = 1, 58; NS). MOUFLON sections hosted fewer

species than DEER or NONE sections (all: F = 3.57; d.f. = 2,

57; P < 0.05; woodland: F = 9.01; d.f. = 2, 57; P < 0.001;

steppe: F = 1.45; d.f. = 2, 57; NS) (Fig. 2). Regarding

management, CLOSED FOREST hosted fewer species than

other management types (F = 11.06; d.f. = 4, 55; P < 0.0001).

More woodland species occurred in STANDARDS and RIDES

than in the remaining three types; whereas steppe species

(F = 4.09; d.f. = 4, 55; P < 0.001) followed a hierarchy

Table 1

Multiple regression models for numbers of all butterfly species, woodland species

Model Model terms

ALL SPECIES Null model

Raw (Kind of game)b � E3 + RE1P

Covariable +NECTAR � SHADE

Controlled +NECTAR � SHADE � E3

WOODLAND spp. Null model

Raw (Kind of game)b + RE1P � RE2 + SPRING

Covariable +NECTAR � SHADE

Controlled +NECTAR � SHADE + RE2 + (Kind of game)b

STEPPE spp. Null model

Raw �E3 + RE1

Covariable +NECTAR

Controlled +NECTAR � E3

ENDANGERED spp. Null model

Raw �E3 + RE1 + SPRING

Covariable +NECTAR

Controlled +NECTAR + (game)c

See Section 2.4 for abbreviations of explanatory terms. Raw models do not consider a

NECTAR, WIND and SHADE, and Controlled models are based on adding explana

individual models; superscript P denotes variable significant in a polynomial forma Comparison of fitted model with null model: ***P < 0.001.b NONE = DEER > MOUFLON.c PRESENT >ABSENT.

(STANDARDS + RIDES) > (COPPICE + CLEARINGS) >CLOSED FOREST. Game ABSENT sections hosted margin-

ally more threatened species than PRESENT sections

(F = 3.72; d.f. = 1, 58; P = 0.06), kind of game exhibited no

effect (F = 2.19; d.f. = 2, 57, NS), and management exhibited a

hierarchy STANDARDS > (CLEARINGS + COPPICE + RI-

RIDES) > CLOSED FOREST (F = 7.66; d.f. = 4, 55; P <0.001).

In the GLM regressions (Table 1), neither section lengths nor

spatial terms exhibited significant effects, leaving nectar and

shade as the only important covariables. They accounted for

more variation for all (79.8%) and steppe (69.7%) butterflies

than for woodland butterflies (45.4%). Still, kind of game was

important for woodland butterflies, whose numbers were lowest

with MOUFLON. Richness of all butterflies decreased with

canopy cover (E3) and displayed a convex response to plant

richness in ground layer (RE1). Richness of woodland

butterflies increased with plant richness (in either ground or

shrub layer), but declined with increasing amount of clearings

adjoining the transect. The only response of steppe species was

a decrease with increasing canopy cover (E3). Finally,

endangered species exhibited a negative correlation with

canopy cover (E3) and a positive correlation with ground plant

richness (RE1). After controlling for nectar, more endangered

butterflies occurred if game was PRESENT.

3.2. Species composition

The first (horizontal) DCA axis (Fig. 3) pointed to a gradient

from woodland butterflies (high ordination scores) to grassland

and steppe species. There were some unexpected patterns, such

as position of P. mnemosyne, an open woodland specialist,

, steppe species and endangered species, observed per transect section

d.f. Devresid qAIC1 F, Pa

1, 60 61.0 61.75

5, 56 20.6 25.07 54.4***

2, 59 12.3 13.62 114.3***

3, 58 10.1 11.50 95.9***

1, 60 61.0 61.96

6, 55 25.9 32.59 12.2***

2, 59 33.3 36.73 24.2***

5, 61 24.4 29.77 16.5***

1, 60 61.0 62.99

2, 59 29.2 32.25 12.5***

1, 60 18.5 19.72 135.8***

2, 59 16.2 17.90 80.1***

1, 60 61.0 61.94

3, 58 26.8 30.65 24.1***

1, 60 17.1 18.27 151.3***

2, 59 15.9 17.60 81.9***

ny covariables, Covariable models are based on forward selection from the terms

tory terms onto the Covariable models. Devresid: residual deviance after fitting

.

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365358

Fig. 3. Unconstrained ordination (DCA) of species at sections of transect

walked in the Milovicky Wood, first and second ordination axes. The ordination

was computed after filtering out spatial autocorrelation effects and average

nectar supply per sections. Eigenvalues of first to fourth axes: 0.14, 0.06, 0.05,

0.04 (sum of all unconstrained eigenvalues: 0.78). Species with weights >1.0

are shown, the symbols show woodland species (triangles), steppe species

(crosses), and species of other habitats, mainly generalists (diamonds).

Fig. 4. Results of the canonical correspondence analyses. Proportions of

variation in distribution of butterflies along a transect across Milovicky Wood,

explained by individual ordination models. All the models were significant at

P < 0.001 level (Monte-Carlo test, 999 permutations). ‘‘Raw models’’ were not

controlled for effects of covariates. ‘‘Spatial models’’ show residual variation

after including latitude, longitude and latitude*longitude interaction of each

separate transect section as covariate terms. ‘‘All covariates’’ models show

residual variation after controlling, in addition to the terms already in spatial

models, for lengths of transect sections and day time, nectar, shade and wind

conditions of each visit.

closely to some grassland species. The second (vertical) axis is

more difficult to interpret, but it seems that species depending

on plants requiring high nitrogen levels (e.g., large nymphalids)

attained higher scores than species requiring nutrient-poor

conditions.

All covariates exhibited significant effects in the CCA

analyses. The strongest effect was that of spatial terms (15.8%

of variation in butterfly records), followed by nectar (10.2%),

weather (8.0%), time of day (5.7%) and section length (3.3%).

The forward selection from covariates returned a model

containing all these terns (44.1% of total variation). However,

all predictors exhibited significant effects (Fig. 4), and their

effects remained significant even after controlling for effects of

covariables. The strongest predictors of butterfly composition

were plant communities, site management and surrounding

management, followed by vegetation covers, plant species

richness, spring herb richness and kind of game.

Kind of game had a stronger effect than just game presence:

DEER sections were more similar to NONE than to

MOUFLON sections (Fig. 5A). Management explained

considerably higher proportion of variation, separating

CLOSED FOREST from any open structures at the first

ordination axis, and COPPICE plus STANDARDS from

CLEARINGS and RIDES at the second axis (Fig. 5B). The

proximity of CLEARINGS and RIDES also appeared in

analysis with surrounding management, in which the first

ordination axis separated closed forest from open structures,

and the second axis pointed to a similarity between

CLEARINGS and CROP FIELDS (Fig. 5C). Adding game-

related (i and ii) variables to models already containing stand

management (iii and iv), and vice versa, revealed significant

independent effects in all combinations tested, identical in

direction to the models not containing the other variables

(Table 2). This applied even in cases when the models

contained covariate terms as well. The same applied for adding

the variables i–iv to models already containing vegetation-

related variables. Therefore, both game keeping and manage-

ment influenced butterfly composition.

4. Discussion

4.1. Game keeping versus management

Game keeping exhibited weaker effects on butterflies than

stand management. Still, the effects of game were statistically

detectable, even after inclusion effects of vegetation or stand

management to the ordinations. Effects of plant composition

were even stronger than effects of management, and although

correlation is not a sign of causation, this hierarchy suggests

that both management and game influence the butterflies via

affecting vegetation (Stewart, 2001; Cote et al., 2004). Some of

the mechanisms causing the influence might be putatively

inferred from the GLM regressions of species richness. For

instance, butterfly richness increased with richness of ground

vegetation, which increases in coppiced stands (Decocq et al.,

2004), but decreases under too high density of game (Chytry

and Danihelka, 1993; Cooke and Farrell, 2001). Richness of

woodland butterflies peaked at intermediate values of ground

plant richness, possibly because too rich ground flora indicates

admixtures of weedy plants.

Practically all butterflies were associated with open

structures, be it standards, coppices, rides or clearings. This

was expectable as only few species tolerate canopy closure in

Central Europe (Benes et al., 2002). Rides and coppices, which

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365 359

Fig. 5. CCA biplots showing positions of individual butterfly species in relation to selected environmental variables. Only butterflies with species with fit values >1

are shown. (A) Kind of game as (categorial predictor), partial ordination after entering spatial covariables into the model. (B) Stand management (categorial

predictor), model not considering spatial covariables. (C) Surrounding management (numeric predictors), model not considering spatial covariables.

Table 2

Effects of adding game and management-related individual environmental variables (=added variable) to ordination models already containing, as covariables, spatial

terms of transect sections, nectar, day time, shade and wind at each visit, plus at least one explanatory variable (=already in v)

Added

variable

Already in v

Game Kind of

game

Management Surrounding

management

Vegetation

covers

Plant spp.

richness

Spring herb

richness

Plant commun.

composition

Ordination axis First All First All First All First All First All First All First All First All

Game – – n.c. n.c. * – * – ** – ** – * – *** –

Kind of game n.c. n.c. – – *** *** ** *** *** *** *** *** *** *** *** ***

Management *** *** *** – – *** *** *** *** *** *** *** *** *** ***

S. managementa *** *** ** *** *** ** * *** ** ** ** *** ** ***

For each model, Monte-Carlo significances of the first and all ordination axes are presented. See Section 2.4 for explanation of the variables tested, n.c.: model not

computable due to lack of variance in predictors. *P < 0.05; **P < 0.01; ***P < 0.001.a Surrounding management.

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365360

are intentionally maintained to benefit the game, are largely

absent from woods grown for timber. In this respect, game

keeping directly supports butterfly diversity. Clearings, which

exist in timber-producing woods as well, represent a different

story, which is evident from the representation of woodland and

steppe butterflies in different types of open structures.

Standards and rides hosted more of the former, in line with

a frequent preference of woodland butterflies for fine-grained

mosaics of sunny and shady conditions (e.g., Sparks et al.,

1994; Valimaki and Itamies, 2005; Cizek and Konvicka, 2005).

For steppe butterflies, standards and rides were as good as

clearings, suggesting that all these structures provide enough

sunlight for open-area butterflies. In addition, rides and

clearings exhibited an affinity to grasslands and crop fields,

whereas coppices and standards exhibited an affinity to closed

forests. Again, these patterns are attributable to vegetation.

Whereas both coppices and standards retain characteristic

woodland plants, grassland plants dominate the rides and

ruderal weeds easily invade the clearings. Indeed, several

butterflies characteristic for farmlands (Pieris rapae, Colias

hyale) were associated with clearings.

Kind of game was more important than game presence, as

minimum butterflies occurred with mouflon. The game is not

native to Central Europe (Hell and Sabados, 1992) and feeds as

a selective herb-preferring grazer. The mouflon sections did not

differ from others in average covers of vegetation layers or in

numbers of plant species, but they contained less nectar

(mouflon: 0.47, deer: 0.72, no game: 1.05; F = 4.97; d.f. = 2,

57; P < 0.05). However, the effect of mouflon remained

significant even in ordinations controlled for such variables as

nectar or plant community composition (Fig. 4), suggesting

more subtle mechanisms. They may include selective grazing

on certain plants. Dolek and Geyer (1997) reported adverse

effects of sheep on sensitive grassland butterflies, and mouflon,

as a closely related animal, might produce similar effects (cf.

Tudor et al., 2004). The mouflon park hosts a higher game

density, which is maintained by providing the animals with

additional fodder. This increases soil nitrogen, promoting

competitively superior plants (Chytry and Danihelka, 1993),

whereas many butterflies depend on poorly competitive stress-

tolerant plants (cf. Dennis et al., 2004; Ockinger et al., 2006).

Therefore, whereas present densities of deer do not harm the

butterflies, and deer keeping even supports them because it

includes intentional maintenance of open structures, the

densities of mouflon are intolerable. We cannot estimate

how much the mouflon should be reduced to eliminate this

harmful effect, but the figure should be lower than current 200

mouflon per 500 ha.

4.2. Steppe butterflies in the wood

A relatively large proportion of butterfly records consisted of

xerophilous species. Some records could have been vagrants

from calcareous steppes of the Palava Hills. Nectar and shade

were particularly important predictors of their species richness,

suggesting that much of their occurrence was driven by short-

term nectar availability. Still, some steppe species occurred in

large numbers (Thymelicus acteon, Minois dryas) and others,

such as burnet moths, are poor dispersers (Menendez et al.,

2002). These two conditions indicate likely breeding in the

wood.

To interpret this, consider that the wide rides support

relatively spacious grassland biotopes, preferred, among others,

by the endangered steppe species T. acteon. Second, whereas

the biotope affinities in Benes et al. (2002) refer to the Czech

Republic as a whole, this study refers to the warmest corner of

the country, in which some xerophilous butterflies shift their

preferences towards cooler and more shady biotopes. Benes

et al. (2003b) and Amiet (2004) document such a shift for

Leptidea sinapis. Finally, the entire distinction between

woodland and steppe butterflies may be an artefact of relatively

recent closure of European forests. Stands resembling coppices

with standards host as many steppe butterflies as rides. Their

affinity to grasslands is clear from ordinations (Fig. 5B), which

document that their butterfly assemblages consist of mixtures of

woodland (e.g. P. mnemosyne) and steppe (Heteropterus

morpheus) species. In a past, when the wood had been actively

coppiced, much more steppe species occurred there

(Appendix A). The persistence of some steppe butterflies

alongside species of open woodlands thus represents a remnant

of a historical landscape with less abrupt distinctions among

biotope types (cf. Rackham, 1998; Konvicka et al., 2006).

4.3. Past and future of Milovicky Wood butterflies

Our results allow a reconstruction of the history of

Milovicky Wood butterflies. The cessation of coppicing in

the 1950s led to canopy closure, which likely restricted open-

forest butterflies to rides, woodland meadows and edges. These

refuges became particularly vulnerable after establishment of

the game parks, as the animals likely preferred them for

stationing, forest meadows were turned to fodder fields. The

first butterflies to disappear included sensitive grassland

specialists, such as some Melitaea spp. (cf. Weiss, 1999), or

species that develop on nutritively attractive legumes, such as L.

morsei, N. sappho or Colias myrmidone (Lorkovic, 1993;

Kudrna and Mayer, 1990; Jutzeler et al., 2000). The

overstocking, peaking in the 1980s, affected even relatively

common species. A notable example is Erebia medusa, a

conspicuous species still occurring in wider surroundings of the

wood, which does not tolerate increased levels of soil nutrients

(Schmitt, 1993). On the other hand, the maintenance of open

structures associated with game keeping supported some open

woodland specialists, such as E. maturna or L. achine, for

longer than in woods managed for timber (Benes et al., 2002).

However, the extent of suitable open structure ultimately

became too limited to ensure long-term survival for many

specialised butterflies. The large-scale rejuvenation cuts,

launched after 2000, came probably too late.

Game keeping thus depleted resources for some species

during a period of high stock densities, but maintained a steady

supply of open structures. Current game densities are much less

harmful, particularly in the deer park. The butterflies also

benefit from the recent rejuvenation cuts that mimic coppicing

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365 361

with standards. This particularly applies to P. mnemosyne,

whose recent population in the wood is largest in the country

(3000 adults in 2006; unpublished data). However, the suitable

conditions are only temporary, as establishment of a high forest

of generative origin remains a long-term management goal;

even the rejuvenation cuts are oriented towards this objective.

This is incompatible with the long-term survival of P.

mnemosyne (Konvicka and Kuras, 1999) and other species

with similar requirements (Warren, 1985, 1991; Greatorex-

Davies et al., 1992; Konvicka et al., 2005). Data from

abandoned coppices from nearby Palava Hills also illustrate

that canopy closure severely depletes ground flora (Hedl,

2003).

Our results illustrate that growing high forest, which did not

exist in the area for the last 1000 years, is incompatible with

preserving the biodiversity of the wood. It is hence not

permissible under the NATURA 2000 status of the area. High

forest is also hardly reconcilable with intensive game keeping,

which increases costs for fencing and protection of nursery

trees. Furthermore, the prospects of growing high-revenue

timber remain questionable, given locally dry climate and

shortages of rainfall. In contrast, coppice management aimed

on fuel wood production (including coppice with standards) is

optimal for butterflies and reconcilable with game keeping.

Vegetative regeneration eliminates costs of tree planting, fresh

coppice provides ideal shelter and deer browse. Therefore,

change of the current policy of growing high forest towards re-

establishment of active coppicing, operated together with game

keeping, is a solution supported by both economic and

conservation considerations (Warren and Key, 1991; Buckley,

1992; Decocq et al., 2005).

From a (butterfly) conservation perspective, a minimum

management goal should be preserving viable populations of

all threatened species still surviving in the wood. To achieve

this, active coppicing should provide a continual supply of

open spaces covering minimally an area of the current

Appendix A

Butterflies (incl. Zygaenidae and S. phegea) recorded in the Milov

transect recording in 2004 and 2005. Nomenclature follows Karsh

Recorded Abbreviation Recent

occurrence

Last record N

Zerynthia polyxena Confirmed 2005

Papilio machaon PapMac Confirmed

Iphiclides podalirius IphPod Confirmed

Parnassius mnemosyne ParMne Confirmed

Aporia crataegi Excluded Prior 1980

Antocharis cardamines AntCar Confirmed

Leptidea sinapis LepSin Confirmed

Leptidea reali Possible Prior 1980 W

Leptidea morsei Excluded 1960s F

Pieris brassicae PieBra Confirmed

Pieris napi PieNap Confirmed

Pieris rapae PieRap Confirmed

Pontia daplidice PonEdu Confirmed

Colias hyale ColHya Confirmed

Colias alfacariensis ColAlf Confirmed 2005

regeneration blocks (ca. 200 ha, now supporting local

population of P. mnemosyne). Of course, the total coppiced

area will have to be three to four times larger, as many of

coppice-thriving butterflies prefer the youngest regrowth

phases (<10 years: cf. Warren, 1987), whereas the historical

coppice cycle had been near 30 years in the area (Vybiral,

2003).

Apart from aesthetic perceptions of forestry planners, the

only objections against coppicing concerns its labour

intensity, and resulting lower revenues compared to high

forest. A combination with hunting revenues may consider-

ably change the balance. There are still unresolved problems,

especially concerning economically profitable and ecologi-

cally sustainable game densities. Some of the problems might

be resolved by establishing enclosures with temporarily

restricted access to animals (Saarinen et al., 2005). Such

enclosures are now in fact provided within the regeneration

blocks, they should function similarly to rotational grazing in

grassland conservation (Kruess and Tscharntke, 2002). In any

case, the case of Milovicky Wood illustrates that a

combination of coppice management and hunting offers an

opportunity for preserving biodiversity of sparse lowland

woodlands of Central Europe.

Acknowledgements

We thank the Zidlochovice forest enterprise, a division of

Czech National Forests, Inc., for permission to work in the

Milovicky Wood, granted to us despite expecting uncomfor-

table results. The following personnel were particularly helpful:

T. Blaha, M. Hrib, J. Joch, J. Vybiral, and first of all, P.

Martinasek. D. Cizkova helped with botanical sampling, J.

Danihelka revised identification of more difficult plants. D.

Hauck, V. Hula, L. Spitzer and P. Vlasanek provided their

recent butterfly records. The study was funded by the Grant

Agency of the Czech Republic (526/04/0417).

icky Wood, Czech Republic, and summary data for quantitative

olt and Razowski (1996).

ote Biotope Conservation

concern

Seen at

transect

Number

seen

w Yes No –

Yes 3

s Yes Yes 9

w Yes Yes 368

s Yes No –

Yes 160

s Yes Yes 74

etter part of the wood No –

ocused search w Yesa No –

Yes 6

w Yes 2064

Yes 324

Yes 6

Yes 32

s No –

Appendix A (Continued )Recorded Abbreviation Recent

occurrence

Last record Note Biotope Conservation

concern

Seen at

transect

Number

seen

Colias erate Confirmed 2004 No –

Colias croceus Confirmed 2004 Migrant No –

Colias myrmidone Excluded Prior 1980 s Yes No –

Gonepteryx rhamni GonRha Confirmed Yes 3

Apatura ilia ApaIli Confirmed w Yes 3

Apatura iris Possible Prior 1980 Elusive w No –

Limenitis camilla Confirmed 2005 w Yes No –

Limenitis populi Possible 1980–1994 Elusive w No –

Neptis sappho Excluded Prior 1950 w Yesa No –

Nymphalis polychloros NymPol Confirmed w Yes 1

Nymphalis xanthomelas Excluded Prior 1950 w Yesa No –

Nymphalis antiopa Confirmed 2005 w No –

Aglais urticae AglUrt Confirmed Yes 11

Vanessa cardui VanCar Confirmed Yes 90

Vanessa atalanta VanAta Confirmed Yes 68

Inachis io InaIo Confirmed Yes 408

Polygonia c-album PolC-al Confirmed w Yes 121

Araschnia levana AraLev Confirmed Yes 357

Euphydryas maturna Excluded Prior 1990 Focused search w Yes No –

Melitaea athalia MelAth Confirmed w Yes 364

Melitaea aurelia Excluded Prior 1980 Focused search s Yes No –

Melitaea phoebe Excluded Prior 1980 Focused search s Yes No –

Melitae didyma Excluded Prior 1980 Focused search s Yes No –

Melitaea trivia Excluded Prior 1980 Focused search s Yesa No –

Melitaea britomartis Excluded Prior 1980 Focused search s Yes No –

Argynnis paphia ArgPap Confirmed w Yes 253

Argynnis adippe Excluded Prior 1980 Focused search w Yes No –

Issoria lathonia IssLat Confirmed Yes 57

Clossiana euphrosyne BolEup Confirmed w Yes Yes 1

Clossiana dia BolDia Confirmed Yes 209

Clossiana selene Excluded 1980–1994 Focused search w No –

Melanargia galathea MelGal Confirmed Yes 2351

Aphantopus hyperanthus AphHyp Confirmed Yes 1731

Maniola jurtina ManJur Confirmed Yes 912

Erebia medusa Excluded Prior 1980 w No –

Coenonympha arcania CoeArc Confirmed w Yes 622

Coenonympha glycerion CoeGly Confirmed Yes 327

Coenonympha pamphilus CoePam Confirmed Yes 251

Lasiommata maera LasMae Confirmed w Yes 195

Lasiommata megaera LasMeg Confirmed Yes 21

Brintesia circe BriCir Confirmed s Yes 294

Hipparchia fagi HipFag Confirmed s Yes Yes 20

Hipparchia semele Excluded Prior 1980 s Yes No –

Minois dryas MinDry Confirmed s Yes Yes 125

Arethusana arethusa AreAre Confirmed s Yes Yes 4

Pararge aegeria ParAeg Confirmed w Yes 97

Lopinga achine Excluded 2001 Focused search w Yes No –

Celastrina argiolus CelArg Confirmed w Yes 45

Lycaena phlaeas LycPhl Confirmed Yes 3

Lycaena dispar LycDis Confirmed Yes 2

Lycaena tityrus LycTit Confirmed 2004 No –

Callophrys rubi CalRub Confirmed w Yes 3

Thecla betulae TheBet Confirmed w Yes 2

Neozephyrus quercus QueQue Confirmed w Yes 361

Satyrium w-album Confirmed 2004 w Yes No –

Satyrium ilicis Possible Prior 1980 Elusive w Yes No –

Satyrium pruni Possible Prior 1980 Elusive s No –

Cupido minimus CupMin Confirmed 2005 s No –

Everes decolorata CupDec Confirmed s Yes 17

Everes argiades CupArg Confirmed s Yes 20

Everes alcetas Excluded Prior 1950 s Yes No –

Glaucopsyche alexis Confirmed 2004 s Yes No –

Plebeius argyrognomon PleArg Confirmed s Yes 13

Plebeius argus Confirmed 2004 s No –

Appendix A (Continued )Recorded Abbreviation Recent

occurrence

Last record Note Biotope Conservation

concern

Seen at

transect

Number

seen

Aricia agestis AriAge Confirmed s Yes 15

Polyommatus icarus PolIca Confirmed Yes 249

Meleageria coridon PolCor Confirmed s Yes 28

Meleageria bellargus PolBel Confirmed 2004 s Yes No –

Meleageria daphnis PolDap Confirmed s Yes Yes 51

Hamearis lucina HamLuc Confirmed w Yes Yes 2

Erynnis tages EryTag Confirmed s Yes 66

Pyrgus malvae PyrMal Confirmed 2004 w No –

Carcharodus alceae CarAlc Confirmed 2005 s Yes No –

Carterocephalus palaemon CarPal Confirmed w Yes 105

Heteropterus morpheus HetMor Confirmed s Yes 88

Thymelicus lineola ThyLin Confirmed Yes 570

Thymelicus sylvestris ThySyl Confirmed w Yes 373

Thymelicus action ThyAct Confirmed s Yes Yes 135

Hesperia comma HesCom Confirmed s Yes Yes 5

Ochlodes venata OchSyl Confirmed Yes 648

Zygaena brizae ZygBri Confirmed s Yes Yes 1

Zygaena osterodensis ZygOst Confirmed w Yes Yes 3

Zygaena loti ZygLot Confirmed Yes 8

Zygaena filipendulae ZygFil Confirmed Yes 6

Zygaena ephialtes ZygEph Confirmed Yes 6

Zygaena angelicae ZygAng Confirmed s Yes 3

Zygaena viciae ZygVic Confirmed Yes 3

Zygaena lonicerae Confirmed 2004 No –

Zygaena carniolica Confirmed 2004 s No –

Syntomis phegea SynPhe Confirmed w Yes 464

Abbreviation: as used in the ordination diagrams.a Species is extinct in the Czech Republic.

Appendix B

The spring aspect survey recorded all dicotyledoneous

plants in bloom by April or earlier (‘‘spring geophytes’’). The

data consisted of 19 species, with mean per section = 4.2 (2.58

S.D.) and range 0–10. The summer aspect survey considered all

higher plants. Some taxonomically difficult flocks of micro-

species (e.g., Taraxacum spp., Rubus spp., Crataegus spp.)

were recorded as aggregates. A total of 295 taxa were recorded.

Split into vegetation layers, the numbers were 18 (E3), 29 (E2)

and 290 (E1); note that some woody species could occur in

several layers. The respective means and ranges per transect

section were 1.6 � 1.59 S.D., 0–6 (E3); 3.5 � 3.38 S.D., 0–11

(E2); 42.7 � 16.3, 8–84 (E1).

The PCA analysis used to characterise plant communities

was based on the summer aspect survey. Covers of all taxa were

expressed as the ordinal values 1–7, taxa recorded in more than

one layer were treated as separate ‘‘species’’ for each layer.

Scaling of the ordination focused on inter-species correlation,

species scores were divided by standard deviations, both

species and samples were centered.

The eigenvlalues of resulting four ordination axes were 0.30.

0.14, 0.08 and 0.06; the respective explained (cumulative)

variances in species data were 30.2%, 44.2%, 52.0% and

58.0%. The first ordination axis pointed to a strong gradient

from sites with high covers of trees and shade-tolerant herbs

(species with highest absolute values of ordination scores: Q.

petrea [E3], Viola mirabilis, Fraxinus angustifolia [E3],

Fallopia dumetorum, Impatiens parviflora, Galium odoratum,

Dactylis polygamma) to sunny sites with light demanding herbs

(Calamagrostis epigeios, Poa angustifolia, Potentilla argentea,

Origanum vulgare, Fragaria vesca, Myosotis arvensis, Achillea

colina). The second axis separated sites with high cover of

shrub layer (A. campestre [E2], Coryllus avellana [E2], Cornus

sanguinea [E2], Crataegus spp. [E2], C. mas [E2], U. laevis

[E2], Rosa canina [E2]), to light-demanding herbs (Hypericum

perforatum, P. angustifolia, O. vulgare, Lactuca seriola,

Dactyllis glomerata, Clinopodium vulgare, Coronilla varia).

References

Absolon, K., 1949. Moravia in Palaeolithic times. Am. J. Archaeol. 53,

19–28.

Amiet, J.L., 2004. Separation des niches ecologiques chez deux especes

jumelles sympatriques de Leptidea (Lepidoptera, Pieridae). Rev. Ecol.

Terre Vie 59, 433–452.

Bakker, E.S., Olff, H., Vandenberghe, C., De Maeyer, K., Smit, R., Gleichman,

J.M., Vera, F.W.M., 2004. Ecological anachronisms in the recruitment of

temperate light-demanding tree species in wooded pastures. J. Appl. Ecol.

41, 571–582.

Benes, J., Konvicka, M., Dvorak, J., Fric, Z., Havelda, Z., Pavlıcko, A., Vrabec,

V., Weidenhoffer, Z., 2002. Butterflies of the Czech Republic: Distribution

and conservation I, II. SOM, Prague.

Benes, J., Kepka, P., Konvicka, M., 2003a. Limestone quarries as refuges for

European xerophilous butterflies. Conserv. Biol. 17, 1058–1069.

Benes, J., Konvicka, M., Vrabec, V., Zamecnik, J., 2003b. Do the sibling species

of small whites, Leptidea sinapis and L. reali (Lepidoptera, Pieridae) differ

in habitat preferences? Biologia 58, 943–951.

Bergman, K.O., 2001. Population dynamics and the importance of habitat

management for conservation of the butterfly Lopinga achine. J. Appl. Ecol.

38, 1303–1313.

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365364

Bergman, K.O., Kindvall, O., 2004. Population viability analysis of the

butterfly Lopinga achine in a changing landscape in Sweden. Ecography

27, 49–58.

Birks, H.J.B., 2005. Mind the gap: how open were European primeval forests?

Trends Ecol. Evol. 20, 154–156.

Bradshaw, R.H.W., Hannon, G.E., Lister, A.M., 2003. A long-term perspective

on ungulate-vegetation interactions. For. Ecol. Manage. 181, 267–280.

Buckley, G.B. (Ed.), 1992. The Ecological Effects of Coppicing. Chapman &

Hall, London.

Chytry, M., Danihelka, J., 1993. Long-term changes in the field layer of oak and

oak-hornbeam forests under the impact of deer and mouflon. Folia Geobot.

Phytotax. 28, 225–245.

Cizek, O., Konvicka, M., 2005. What is a patch in a dynamic metapopulation?

Mobility of an endangered woodland butterfly Euphydryas maturna.

Ecography 28, 791–800.

Cooke, A.S., Farrell, L., 2001. Impact of muntjac deer (Muntiacus reevesi) at

Monks Wood National Nature Reserve, Cambridgeshire, eastern England.

Forestry 74, 241–250.

Cote, S.D., Rooney, T.P., Tremblay, J.P., Dussault, C., Waller, D.M., 2004.

Ecological impacts of deer overabundance. Ann. Rev. Ecol. Evol. Syst. 35,

113–147.

Decocq, G., Aubert, M., Dupont, F., Alard, D., Saguez, R., Wattez-Franger, A.,

De Foucault, B., Delelis-Dusollier, A., Bardat, J., 2004. Plant diversity in a

managed temperate deciduous forest: understorey response to two silvi-

cultural systems. J. Appl. Ecol. 41, 1065–1079.

Decocq, G., Aubert, M., Dupont, F., Bardat, J., Wattez-Franger, A., Saguez, R.,

de Foucault, B., Alard, D., Delelis-Dusollier, A., 2005. Silviculture-driven

vegetation change in a European temperate deciduous forest. Ann. For. Sci.

62, 313–323.

Dennis, R.L.H., Hodgson, J.G., Grenyer, R., Shreeve, T.G., Roy, D.B., 2004.

Host plants and butterfly biology. Do host-plant strategies drive butterfly

status? Ecol. Entomol. 29, 12–26.

Dolek, M., Geyer, A., 1997. Influence of management on butterflies of rare

grassland ecosystems in Germany. J. Insect Conserv. 1, 125–130.

Feber, R.E., Brereton, T.M., Warren, M.S., Oates, M., 2001. The impacts of deer

on woodland butterflies: the good, the bad and the complex. Forestry 74,

271–276.

Freese, A., Benes, J., Bolz, R., Cizek, O., Dolek, M., Geyer, A., Gros, P.,

Konvicka, M., Liegl, A., Stettmer, C., 2006. Habitat use of the endangered

butterfly Euphydryas maturna and forestry in Central Europe. Anim.

Conserv. 9, 388–397.

Fuller, R.J., Gill, R.M.A., 2001. Ecological impacts of increasing numbers of

deer in British woodland. Forestry 74, 193–199.

Greatorex-Davies, J.N., Hall, M.L., Marrs, R.H., 1992. The conservation of

pearl-bordered fritillary butterfly (Bolororia euphrosyne L.): preliminary

studies on the creation and management of glades in conifer plantations.

For. Ecol. Manage. 53, 1–14.

Greatorex-Davies, J.N., Sparks, T.H., Jall, M.L., Marrs, R.H., 1993. The

influence of shade on butterflies on ridges of coniferised lowland woods

in southern England and implications for conservation management. Biol.

Conserv. 63, 31–41.

Hansson, L., 2001. Traditional management of forests: plant and bird commu-

nity responses to alternative restoration of oak-hazel woodland in Sweden.

Biodiv. Conserv. 10, 1865–1873.

Hedl, R., 2003. Lesni vegetace NPR Devin (CHKO a BR Palava) po 50 letech

samovolneho vyvoje a alternativy budouciho hospodareni. In: Karas, J.

(Ed.), Vliv Hospodarskych Zasahu a Spontanni Dynamiky Porostu na stav

Lesnich Ekosystemu. Sbornik Prispevku z Konference, CZU, Kostelec nad

Cernymi lesy, unpaginated (in Czech).

Hell, P., Sabados, K., 1992. Das Muffelwild in der Tschechoslowakei. Zeits-

chrift fur Jagdwissenschaft 38, 9–15.

Homolka, M., Heroldova, M., 2003. Impact of large herbivores on mountain

forest stands in the Beskydy Mountains. For. Ecol. Manage. 181, 119–

129.

Joys, A.C., Fuller, R.J., Dolman, P.M., 2004. Influences of deer browsing,

coppice history, and standard trees on the growth and development of

vegetation structure in coppiced woods in lowland England. For. Ecol.

Manage. 202, 23–37.

Jutzeler, D., Hoettinger, H., Malicky, M., Rebeusek, F., Sala, G., Verovnik, R.,

2000. Biology of Neptis sappho (Pallas, 1771) based on the monograph by

Timpe ( Timpe (1993) and its actual distribution and conservation status in

Austria, Italy and Slovenia (Lepidoptera: Nymphalidae). Linneana Belgica

17, 315–330.

Karsholt, O., Razowski, J., 1996. The Lepidoptera of Europe, Distributional

Checklist. Apollo Books, Stenstrup.

Konvicka, M., Kuras, T., 1999. Population structure, behaviour and selection

of oviposition sites of an endangered butterfly, Parnassius mnemosyne,

in Litovelske Pomoravı Czech Republic. J. Insect Conserv. 3, 211–

223.

Konvicka, M., Cizek, O., Filipova, L., Fric, Z., Benes, J., Krupka, M.,

Zamecnik, J., Dockalova, Z., 2005. For whom the bells toll: demography

of the last population of the butterfly Euphydryas maturna in the Czech

Republic. Biologia 60, 551–557.

Konvicka, M., Vlasanek, P., Hauck. D. 2006. Absence of forest mantles creates

ecological traps for Parnassius mnemosyne (Lepidoptera, Papilionidae).

Nota Lepid., in press.

Kruess, A., Tscharntke, T., 2002. Grazing intensity and the diversity of grass-

hoppers, butterflies, and trap-nesting bees and wasps. Conserv. Biol. 16,

1570–1580.

Kudrna, O., Mayer, L., 1990. Grundlagen zu einem Artenhifsprogramm fur

Colias myrmidone (Esper, 1780) in Bayern. Oedippus 1, 1–46.

Liegl, A., Dolek, M., 2006. Conservation of coppice with standards for canopy

arthropods. In: Floren, A., Schmidl, J. (Eds.). Canopy Arthropod Research

in Central Europe: Basic and Applied Studies from the High Frontier.

Nurnberg, Bioform, in press.

Lorkovic, Z., 1993. Ecological association of Leptidea morsei major (Grund

1905) (Lepidoptera, Pieridae) with the oak forest Lathyro-quercetum

peetraeae HR.-T. 1957 in Croatia. Period. Biol. Zagreb 95, 455–457.

MathSoft, 1999. S-Plus 2000. Guide to Statistics, Volume 1. Data Analysis

Products Division. MathSoft Inc., Seattle, WA.

Menendez, R., Gutierrez, D., Thomas, C.D., 2002. Migration and Allee effects

in the six-spot burnet moth Zygaena filipendulae. Ecol. Entomol. 27, 317–

325.

Ockinger, E., Hammarstedt, O., Nilsson, S.G., Smith, H.G., 2006. The relation-

ship between local extinctions of grassland butterflies and increased soil

nitrogen levels. Biol. Conserv. 128, 564–573.

Pollard, E., 1977. A method for assessing changes in the abundance of

butterflies. Biol. Conserv. 12, 115–134.

Pollard, E., Cooke, A.S., 1994. Impact of muntjac deer Muntiacus reevesi on

egg-laying sites of the white admiral butterfly Ladoga camilla in a Cam-

bridgeshire wood. Biol. Conserv. 70, 189–191.

Pollard, E., Woiwod, I.P., Greatorex-Davies, J.N., Yates, T.J., Welch, R.C.,

1998. The spread of coarse grasses and changes in numbers of lepidoptera in

a woodland nature reserve. Biol. Conserv. 84, 17–24.

Povolny, D., Gregor, F., 1946. Vretenusky (Zygaena Fab.) v zemi Moravsko-

slezske Entomol. listy (Suppl. 12), 1–100 (in Czech).

Rackham, O., 1998. Savanna in Europe. In: Kirby, K.J., Watkins, C. (Eds.), The

Ecological History of European Forests. CAB International, Wallingford,

pp. 1–24.

Saarinen, K., Jantunen, J., Valtonen, A., 2005. Resumed forest grazing restored

a population of Euphydryas aurinia (Lepidoptera: Nymphalidae) in SE

Finland. Eur. J. Entomol. 102, 683–690.

Schmitt, T., 1993. Biotopanspruche von Erebia medusa brigobanna Fruhstorfer,

1917 (Rundaugen-Mohrenfalter) im Nordsaarland (Lepidoptera, Nympha-

lidae, Satyrinae). Atalanta 24, 33–56.

Skala, H., 1912–1913. Die Lepidopterenfauna Mahrens I. Verh. naturforsch.

Ver. Brunn 50, 63–241.

Sparks, T.H., Porter, K., Greatorex-Davies, J.N., Hall, M.L., Marrs, R.H.,

1994. The choice of oviposition sites in woodland by the Duke of

Burgundy butterfly Hamearis lucina in England. Biol. Conserv. 70,

257–264.

Stewart, A.J.A., 2001. The impact of deer on lowland woodland invertebrates: a

review of the evidence and priorities for future research. Forestry 74, 259–

270.

ter Braak, C.J.F., Smilauer, P., 1998. CANOCO Reference Manual and User’s

Guide to Canoco for Windows. Software for Canonical Community

J. Benes et al. / Forest Ecology and Management 237 (2006) 353–365 365

Ordination (version 4). Centre for Biometry Wageningen (Wageningen, NL)

and Microcomputer Power, Ithaca, NY, USA.

Tudor, O., Dennis, R.L.H., Greatorex-Davies, J.N., Sparks, T.H., 2004.

Flower preferences of woodland butterflies in the UK: nectaring spe-

cialists are species of conservation concern. Biol. Conserv. 119, 397–

403.

Valimaki, P., Itamies, J., 2005. Effects of canopy coverage on the immature

stages of the Clouded Apollo butterfly [Parnassius mnemosyne (L.)]

with observations on larval behaviour. Entomol. Fennica 16, 117–

123.

Van Swaay, C.A.M., Warren, M.S., 1999. Red Data Book of European

Butterflies (Rhopalocera). Nature and Environment Series No. 99. Council

of Europe, Strasbourg.

Vera, F.W.M., 2000. Grazing Ecology and Forest History. CAB International,

Wallingford.

Vrabec, V., Lastuvka, Z., Benes, J, Sumpich, J., Konvicka, M., Fric, Z., Hrncir,

J., Matous, J., Marek, S., Kuras, T., Hula, V., Herman, P., 2006. Lepidoptera

(motyli). In: Farkac, J., Kral, D., Skorpik, M. (Eds.), Cerveny seznam

bezobratlych zivocichu. Priroda, Praha, pp. 172–236 (in Czech, English

summary).

Vybiral, J., 2003. Lesy a myslivost v Chranene krajinne oblasti Palava. In:

Danihelka, J. (Ed.), Palava na Prahu Tretiho Tisicileti. CHKO Palava,

Mikulov, (in Czech), pp. 105–112.

Wahlberg, N., Klemetti, T., Hanski, I., 2002. Dynamic populations in a dynamic

landscape: the metapopulation structure of the marsh fritillary butterfly.

Ecography 25, 224–232.

Warren, M.S., 1985. The influence of shade on butterfly numbers in woodland

rides, with special reference to the Wood White Leptidea sinapis. Biol.

Conserv. 33, 147–164.

Warren, M.S., 1987. The ecology and conservation of the heath fritillary,

Mellicta athalia. III. Population dynamics and the effect of habitat manage-

ment. J. Appl. Ecol. 24, 499–513.

Warren, M.S., 1991. The successful conservation of an endangered species, the

Heath fritillary butterfly Mellicta athalia in Britain. Biol. Conserv. 55, 37–56.

Warren, M.S., Key, R.S., 1991. Woodlands: past, present and potential for

insects. In: Collins, N.M., Thomas, J.A. (Eds.), The Conservation of Insects

and Their Habitats. Academic Press, London, pp. 155–212.

Weiss, S.B., 1999. Cars, cows, and checkerspot butterflies: nitrogen deposition

and management of nutrient-poor grasslands for a threatened species.

Conserv. Biol. 13, 1476–1486.