Ann. For. Res.

57(2): 215-231, 2014 ANNALS OF FOREST RESEARCH

DOI: 10.15287/afr.2014.172 www.afrjournal.org

Faunal diversity of Fagus sylvatica forests: A regional

and European perspective based on three indicator
groups

H. Walentowski, S. Müller-Kroehling, E. Bergmeier, M. Bernhardt-Römermann, M.M.

Gossner, A. Reif, E.-D. Schulze, H. Bußler, C. Strätz, W. Adelmann

Walentowski H., Müller-Kroehling S., Bergmeier E., Bernhardt-Römermann, M.,

Gossner M.M., Reif A., Schulze E.-D., Bußler H., Strätz C., Adelmann W., 2014.
Faunal diversity of Fagus sylvatica forests: A regional and European perspective
based on three indicator groups. Ann. For. Res. 57(2): 215-231, 2014.

Abstract. While the postglacial history of European beech (Fagus sylvatica)

and the plant species composition of beech forests in Central Europe are
fairly well understood, the faunal biodiversity has been less well investiga-
ted. We studied three groups of mostly sedentary organisms in beech forest
at regional and European scales by combining field studies with a compila-
tion of existing literature and expert knowledge. Specifically, we examined
the relationship between host tree genera and saproxylic beetles, and the
diversity and composition of forest ground-dwelling molluscs and ground
beetles in relation to the abundance of beech. At a West Central European
scale (Germany), where beech has a “young” ecological and biogeographi-
cal history, we found 48 primeval forest relict species of saproxylic beetles
associated with beech, 124 ground beetles and 91 molluscs inhabiting beech
forest, yet none exclusive of West Central European beech forests. High lev-
els of faunal similarity between beech and other woodland trees suggested
that many of the beech forest dwelling species are euryoecious and likely to
originate from mid-Holocene mixed broadleaf forests. Beech forests of the
mountain ranges in Southern and East Central Europe, which are ecologi-
cally and biogeographically “old”, were found to harbour distinct species
assemblages, including higher contingents of saproxylic beetles and cara-
bids in the Carpathians and narrow-range endemic gastropods of Carpathian
broadleaf forest.The observed biodiversity patterns suggest differentiated
conservation priorities in “young” and “old” European beech forest regions.
Keywords beech forests, faunal biodiversity, Carabidae, conservation pri-
orities, ecological history, habitat continuity, Mollusca, primeval forest rel-
icts, saproxylic beetles.

Authors. Helge Walentowski ([email protected]), S. Müller-

Kroehling, H. Bußler - Bavarian State Institute of Forestry, Freising, Germany;
E. Bergmeier - Department of Vegetation & Phytodiversity Analysis, Univer-
sity of Göttingen, Germany; M. Bernhardt-Römermann - Institute of Ecology,
Friedrich Schiller University Jena, Germany; M.M. Gossner - Terrestrial Eco-

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Ann. For. Res. 57(2): 215-231, 2014 Research article

logy Research Group, Department of Ecology and Ecosystem Management,

Center for Food and Life Sciences Weihenstephan, Munich University of Tech-
nology, Germany; A. Reif - Department of Vegetation Science and Site Classifi-
cation, University of Freiburg, Germany; E.-D. Schulze - Max-Planck-Institute
for Biogeochemistry, Jena, Germany; Strätz C. - Office for Ecological Studies,
Bayreuth, Germany; Adelmann W. - Bavarian Academy for Nature Conserva-
tion and Landscape Management, Germany.
Manuscript received May 01, 2014; revised August 08, 2014; accepted August 13,
2014; online first August 22, 2014.

Introduction schke & Bohn 2004). Regional monographs

emphasise both abiotic factors and the role of
The biogeographic patterns we observe today humans in shaping beech woodlands and their
are highly influenced by current and past envi- distribution (e.g., Horvat et al. 1974, Dierßen
ronmental conditions, human impact, soil de- 1996, Ellenberg & Leuschner 2010). Willner et
velopment and biological evolution (Fang and al. (2009) showed that the distribution of many
Lechowicz 2006). European beech (Fagus syl- range-restricted beech forest understorey plants
vatica L.) forest development in West, Central may be explained by dispersal limitations rath-
and Northern Europe has been very different, er than by their environmental requirements. In
compared to South and South-Eastern Europe contrast, the faunal species diversity and its de-
(Magri et al. 2006, Giesecke et al. 2007). For velopment in European beech forests, beyond
example, Illyric beech forests in the western local studies, are far less well known.
part of the Balkan Peninsula occur at least since In the past, animal species either followed
early postglacial times (10300–8800 years BP), beech from their refugia or were already pres-
and they are particularly diverse in forest plant ent in the region and colonised the expanding
species (Horvat et al. 1974). In contrast, North beech forests. In the first case, at least some
of the Alps beech appeared first approximately specialised beech forest dwellers may be ex-
8000 years BP (Tinner & Lotter 2006). In the pected, while in the second case, exclusive
period of Subboreal (4500–3000 years BP), the beech specialists may be lacking unless there
mixed mesophytic upland and lowland forests is evidence of recent speciation. To test for
were subjected to several waves of beech inva- potential mechanisms explaining faunal spe-
sion (Pott 1989, Pott 1997), but it was not until cies composition of European beech forests,
the Subatlantic period (3000–1000 years BP) the distinctiveness of species pools rather than
that the composition finally shifted towards species numbers is to be analysed. Habitat-re-
nearly pure beech forests. lated species pools may comprise: (i) ubiqui-
Beech forests provide habitat conditions tous, (ii) range-restricted but euryoecious, (iii)
different from those of other broadleaf forest stenoecious species. The proportion of each in
types, characterised by deep shade, leaf litter a biotope or region may serve as an indicator of
durability and accumulation, and a rather spe- the ecological history and distinctiveness.
cies-poor shrub and moss layer (Delagrange et The aim of our study was to revealing pat-
al. 2006, Valladares & Niinemets 2008, Brunet terns of ecological distinctiveness of Euro-
et al. 2010). The plant species composition of pean beech forest using three indicator animal
European beech forests and their geographical groups with sufficient data available, saprox-
and ecological differentiation are well known ylic beetles, woodland ground beetles (Cara-
(e.g., Dierschke 1990, Diekmann et al. 1999, bidae), and woodland molluscs. The selected
Bohn et al. 2000/2003, Willner 2002, Dier- groups differ in their association to specific
216
Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

forest structures (e.g. deadwood), and they particular tree genus.

represent different trophic levels, feeding hab- We considered mesoscale (Germany or part
its and dispersal potential. thereof) and macroscale (multiple European
Saproxylic beetles play an important role in countries) faunal diversity patterns and infer
wood decomposition and nutrient cycling in that there are different implications for con-
forest ecosystems (Nieto & Alexander 2010). servation management in ecologically and bio-
They form a taxonomically and, apart from geographically “young” and “old” beech forest
their dependence on dead or dying wood, eco- regions.
logically heterogenous group. Generally, many
saproxylic beetles are associated with specific
properties of old growth stands such as stand Materials and methods
continuity and the occurrence of ancient trees
(Nilsson & Baranowsky 1993, 1994; Müller et Species groups investigated
al. 2012).
Ground beetles (Carabidae) are important (i) As the range of host trees of Germany`s
predators in forest ecosystems (Lovei & Sun- 1378 saproxylic beetle species (Schmidl &
derland 1996) and include poor, medium and Bußler 2004) is insufficiently known and even
well-dispersing species, with only the latter less so for Europe as a whole, we selected a
exhibiting flight capability (Riley & Browne subset of saproxylic beetle species for which
2011). Many species are restricted to particu- reliable host tree data in Germany and Europe
lar habitats (Gutiérrez et al. 2003). Due to their exists. Müller et al. (2005) provided a list of
species diversity and dispersal limitations, 115 central European saproxylic beetles of
and due to their important role in ecosystems, primeval forest (PFR hereafter, “Urwaldre-
ground beetle assemblages reflect well the liktarten” in germ.), among them some of the
ecological history and conservation value of most endangered European species (Jansson
forest habitats. Land snails (Gastropoda) and 2009). We compiled the published tree genus
well preserved fossil snail shells from lime- records of the 115 PFR species from Germany,
stone and Holocene calcareous tufa sediments Austria, France, Finland, Sweden and Roma-
have been widely used as indicators for (pal- nia, supplemented by our additional field notes
aeo-)environmental conditions and changes from Romania, Poland, Finland and Germany
(Gedda 2001, Meyrick 2003, Gedda 2006, (see Supporting Information I for details on the
Mitrović 2007). A special feature of this group underlying literature).
is its limited dispersal potential. (ii) For ground beetles, we used a dataset
We hypothesise that in West Central Europe comprising 622 plots, with 415 in Germany
where beech forests are ecologically and bio- and 207 elsewhere in Central, South Eastern
geographically “young”, (i) the period of beech and Eastern Europe (see Table 1 and Support-
dominance has been too short to beget beech ing Information II for details about the studies
forest specialists and hence, if there are any included). A total of 381 plots (61%) were re-
in west central Europe, they most likely fol- lated to beech forests, while 241 sample plots
lowed Fagus during its postglacial expansion; were in ravine forests and in various mostly
(ii) relict species of primeval forest are hardly near-natural oak, lime, pine and spruce forests
beech forest specialists, but rather adapted to (Table 1). The dataset includes log transformed
broadleaf forest in general; (iii) structural and abundance data of ground beetle assemblages
tree species diversity of woodlands are crucial per plot and among other information, the
for tree and ground dwelling faunal diversity abundance of beech in the tree layer scaled at
and more important than the presence of any 0 < 1% cover, 1 < 30%, 2 < 50%, 3 < 70%, 4 <

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 Ann. For. Res. 57(2): 215-231, 2014 Research article

90%, 5 > 90%.

Total

381
56

43
32
622
110
(iii) In the absence of comprehensive
mollusc data for Europe or any larger

(Ukraine)
Transcar-
Table 1 Allocation of all 622 ground beetle (Carabidae) sample plots to habitat type and region. Beech forest plots are represented by 381 stands

area (but see Čiliak & Šteffek 2011 for

pathian
Eastern Continental

plains
the Slovak Republic) we established a
mollusc (land snails, slugs and freshwa-

5
ter molluscs) dataset of habitat-related
plain,Tran-
Hungarian

sylvanian

species abundance data from 70 forest

plateau
Plains

reserves in Bavaria, Southern Germany,

representing most natural forest types of
9
2
34

45
this area (Bohn 2000/2003, Walentowski
and Sudeten
Carpathian

subdivided
Mountains

et al. 2001, Walentowski et al. 2013).

We combined that with environmental
Not

data made available through the Forest

61

2
15

78 Reserve Database of the Bavarian State

Alpine- Western
Adriatic Balkans
(Illyric)
western Balkans)

Institute of Forestry (see Supporting In-

(southern Alps,

formation III for details). The mollusc

Southeastern

6
1
1

8
Mountains

dataset is considered representative of

the Central European uplands including
region

parts of the Northern Limestone Alps.

4
2
5

11
Eastern

Studied forest habitat types (ground

Alps

30
5
1
2
9
47

dwellers)
Northern (and
Central) Alps

and Alpine
Swiss Jura
Alps (incl.

foreland)
Western

For the forest ground-dwelling groups

(ground beetles and molluscs) we distin-
9

13

guished six forest habitat types: beech,

(Hercynian-
Central European uplands

mixed montane (Fagus, Abies, Picea),

Polonic)
uplands
Eastern

oak and oak-lime-hornbeam, mixed

ravine and slope woods (Tilia, Ulmus,
170
36
9
26
9
250

Fraxinus, Acer), (Semi)natural pine

(Subatlantic)

woods (Pinus sylvestris), (Semi)natural

uplands and
scarplands

spruce forest (Picea abies). Regarding

Western

molluscs, riparian and other wetland

73
10
39

4
126

forests (Populus, Alnus, Fraxinus) were

included, and lowland to submontane
(Poland)
Baltic

beech forests (often co-dominated by

15

10
34

beech and oak) were separated from

montane forests (often co-dominated
Atlantic)
German

(if not

by beech, fir and spruce). In contrast,

North
Costal lowlands

Plain

for the carabid data set, wetland forests

1

were disregarded as they showed little

Atlantic

of Ger-
region

relationship to beech forests at the Eu-

many

ropean scale, and lowland beech forests

Beech 3

were merged with montane mixed beech

Habitat

Ravine

Spruce
Mixed

forests. Further, the carabid beetle data

Total
Pine
type

oak

218
Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

were related to different levels of forest utili- DCA scores with environmental factors (for
sation intensity: 0-virgin forest, 1- semi-virgin molluscs: mean annual precipitation and tem-
forests (cut in the past but virgin forest struc- perature, elevation, forest type, soil base sta-
tures intact), 2-bordering virgin forest, 3-old tus, in 3 classes - low, medium, high, surface
reserve (not cut in more than 50 years, siz- water, rocks, proportion of beech, for ground
able), 4-young reserve (not cut in less than 50 beetles: forest type, land use intensity, in 4
years, mostly small), 5-managed forest or not classes - none to very intensive, the propor-
cut in less than 20 years. All mollusc data fall tion of beech). To reveal differences between
into category 4. geographic ranges, analyses for saproxylic and
ground beetles were performed separately for
Geographic range, spatial scales and data the entire dataset (Europe) and for selected re-
compilation gions. All statistical analyses were performed
using R 2.14 (R Development Core Team
Data was analysed on two spatial levels, (i) 2011), with the additional packages vegan
mesoscale (Germany for saproxylic and ground (Oksanen et al. 2011) and fpc (Hennig 2010).
beetles and Bavaria for molluscs), assumed to
be representative for west central Europe and
(ii) macroscale (considering saproxylic beetles Results
and carabids), comprising European sites with
short or long distances to glacial broadleaf for- Mesoscale diversity
est refuge areas. To categorise the macroscale
data biogeographically (and along with differ- Fourty-eight (42%) of the 115 PFR saproxy-
ent degrees of forest use), we distinguished six lic beetle species recorded in Germany were
regions: Atlantic and Baltic coastal lowlands, found on beech at least once. For comparison,
Central European uplands, Northern and Cen- 66 PFR species (57%) were found on oak,
tral Alps, Southern Alps and western Balkans, much less on Populus (21; 18%), Salix (19;
Carpathian and Sudeten Mountains, Eastern 17%), Picea (18; 16%), Tilia (15; 13%) and
continental plains (Table 1). Ulmus (14; 12%). The cluster analysis revealed
eight clusters of host trees associated with bee-
Statistical analyses tle assemblages (Fig. 1). Thirteen PFR species
were found to characterise the various groups
We used cluster analysis (Ward’s method with of host tree genera (Table 2).
binary Bray-Curtis distances) to detect asso- Cluster 5 includes beech (Fagus) and the
ciations of saproxylic beetle assemblages (of deciduous tree genera Quercus (most similar
species recorded at least once on beech) with to beech), Ulmus, Populus and Salix. Together
host tree species. The appropriate number of with Carpinus and Acer (cluster 4), being most
clusters was identified using bootstrapping similar in species composition to cluster 5,
methods (Hennig 2010). Canonical Corre- those tree genera are characteristic for lowland
spondence Analysis (CCA) with beech cover and lower upland forest, including both zonal
as the constraining factor linked ground beetle and azonal pioneer woodlands. The next most
and mollusc communities to the abundance of similar clusters (6, 7 and 8) included a range
European beech. Detrended Correspondence of broadleaf trees that represent more or less
Analysis (DCA) was used to identify environ- warm sites in the nemoral forest and cultural
mental factors related to the differences found landscape: cluster 6 - Platanus (introduced),
in the mollusc and ground beetle datasets, re- Pyrus and Malus (native and cultivated), cluster
spectively. Subsequently, we correlated the 7 - Aesculus (introduced), Fraxinus and Tilia,

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Figure 1 Clusters of host tree genera of 48 beech exploiting primeval forest relict
species of saproxylic beetles at the mesoscale (Germany)

Table 2 Saproxylic beetle species indicating groups of host tree genera in Germany (threshold indicator
value [IV] > 40%). Cluster numbers correspond to Fig. 1
cluster IV p - value
Ceruchus chrysomelinus 3 41.1 0.010
Nematodes filum 4 71.4 0.011
Dicerca berolinensis 4 71.4 0.011
Allecula rhenana 4 62.5 0.005
Euryusa coarctata 5 80.0 0.003
Ampedus elegantulus 5 80.0 0.004
Megapenthes lugens 5 60.0 0.001
Ischnodes sanguinicollis 5 50.0 0.003
Neatus picipes 5 46.0 0.001
Elater ferrugineus 5 42.9 0.001
Crepidiphorus mutilatus 7 55.6 0.001
Abraeus parvulus 7 52.6 0.012
Teretrius fabricii 8 55.6 0.004

cluster 8 - Betula and Pinus. Less accordance cea (cluster 3). The PFR species of these gen-
in saproxylic beetle assemblages with beech era inhabit cool mountainous or wet forests.
and oak was observed for Alnus, Abies and Pi- The most pronounced dissimilarities (clusters

220
Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

1 and 2) to beech and oak were observed for ciduous forest (Fig. 2). All carabid assemblag-
Corylus, Juglans (introduced), Prunus (native es found in deciduous forest types overlapped
and cultivated) and Sorbus, presumably due to to a large extent. Beech forest carabid fauna
their different, often shrubby habit, low maxi- overlapped particularly with that of ravine for-
mum age, the resulting lack of deadwood and ests and less so with that of oak forests.
rarity of ancient trees. All saproxylic beetle A total of 181 mollusc species (five of them
PFR species found on beech wood occured on with data deficiencies and thus excluded from
a range of host tree species, although some pre- the DCA analysis) was observed in the 70 Ba-
fer specific groups of host trees. While none is varian forest nature reserves, including land
entirely restricted to beech, six out of 48 beech snails (123 species), slugs (23), freshwater
exploiting PFR species were exclusively as- snails (26) and bivalves (9). The species com-
sociated with the host tree genera of cluster 5 position was strongly depending on tempera-
(Table 2). ture (correlation with first axis: r = 0.5, t =
Among the 191 ground beetle (carabid) spe- 4.79, df = 68, p < 0.001) and correspondingly
cies sampled across the forest types of Germa- on elevation (correlation with first axis: r = -
ny, 124 species (66%) were observed in beech 0.43, t = -3.96, df = 68, p < 0.001), further on
forests, but none of these species were exclu- base content (correlation with first axis: r = -
sive of beech forests. While Pterostichus bur- 0.58, t = 5.83, df = 68, p < 0.001; for second
meisteri did show a significant preference for axis: r = -0.8 t = 11.09, df = 68, p < 0.001)
beech forests (r = 0.60, t = 15.47, df = 426, p < (Fig. 3). The first axis of the DCA showed a
0.001), it has been recorded in ravine forests, gradient from warm floodplain sites to cool
too. The carabid composition of semi-natural montane to subalpine levels of the Alps. The
conifer forests (subalpine spruce forests, pine- second axis showed a gradient from acid-oli-
woods) showed little similarity to that of de- gotrophic sites to eutrophic sites. Assemblages

Figure 2 Ordination diagram (DCA) of the ground beetle assemblages of different forest types in Germany
(from 428 sample plots and 191 species). Axis 1: eigenvalue 0.46, length of gradient 5.09. Axis 2:
eigenvalue 0.29, length of gradient 4.05.
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Ann. For. Res. 57(2): 215-231, 2014 Research article

Figure 3 Ordination diagram (DCA) of the mollusc assemblages (176 species) of different forest types in 70
Bavarian forest reserves. Axis 1: eigenvalue 0.24, length of gradient 3.07. Axis 2: eigenvalue 0.19,
length of gradient 2.71, environmental vectors passively plotted.

of molluscs inhabiting beech forests (91 spe- cies; 70%), while other host tree genera were
cies) showed a scattered distribution in the or- less represented (Populus, 37 species; Picea,
dination plot with similarities to assemblages 29; Abies, 26; Pinus, 25; Salix, 24; Tilia, 24).
of various forest types, particularly of mixed All host tree genera had more associated PFR
oak woodlands and ravine forests. We did not saproxylic beetle species at a European scale
find positive correlations of mollusc species than at a west central European scale.
specifically with beech. A negative correlation The similarities regarding beetle assemblag-
(not shown) between beech dominance and es (Fig. 4) and the correlations between spe-
molluscs of wet forest (Zonitoides, Oxyloma, cies occurrences and host tree clusters (Table
Succinea) and riparian forest (Trochulus) was 3) were similar to those observed at the mes-
to be expected. oscale. Two high-level groupings were distin-
guished: the first group (combining clusters 1
Macroscale diversity to 5) mainly included tree genera of the south-
nemoral transition zone with a more southern
At a European scale, a total of 63 (55%) of the (sub-Mediterranean) or lowland distribution.
115 PFR saproxylic beetles were found to be The second group (clusters 6 to 9) included
exploiters of beech wood, thus 15 more spe- tree genera associated with temperate and
cies than at a central European level. This was cool-temperate zones (nemoral, boreo-nemor-
not merely an effect of the larger area, since al) as well as wetland, pioneer and montane-
also in a single region such as southern Ro- prealpine sites.
mania a higher number of PFR saproxylic Beech forests carabid assemblages differed
beetles associated with beech were observed considerably among regions (Fig. 5). Distinct
(10 species). As for Germany, PFR on beech assemblages were identified in the Carpathian
were outnumbered by those on oak (80 spe- and Sudeten high mountain ranges and in the
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Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

Figure 4 Clusters of host tree genera of 63 beech exploiting primeval forest relict species of saproxylic bee-
tles at the European scale (Germany, Austria, France, Finland, Sweden and Romania)

Table 3 Saproxylic beetle species indicating groups of host tree genera in Europe (threshold indicator
value IV > 40 %). Cluster numbers correspond to Fig. 4
cluster IV p - value
Allecula rhenana 4 62.5 0.003
Nematodes filum 4 48.4 0.026
Mycetoma suturale 6 83.3 0.011
Lacon lepidopterus 6 71.4 0.009
Ipidia binotata 6 48.4 0.028
Ampedus elegantulus 6 43.5 0.017
Abraeus parvulus 8 57.7 0.005
Crepidiphorus mutilatus 8 55.6 0.002
Euryusa coarctata 9 80.0 0.005
Triplax collaris 9 80.0 0.004
Epierus comptus 9 60.0 0.049
Megapenthes lugens 9 60.0 0.001
Ischnodes sanguinicollis 9 54.6 0.001
Neatus picipes 9 46.2 0.002
Elater ferrugineus 9 42.9 0.001
Thoracophorus corticinus 9 40.0 0.001

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Ann. For. Res. 57(2): 215-231, 2014 Research article

Figure 5 Ordination diagram (DCA) of the ground beetle assemblages of beech forests in various European
regions derived from 381 sample plots and 199 species. Axis 1: eigenvalue 0.41, length of gradient
4.62. Axis 2: eigenvalue 0.26, length of gradient 3.31.

Figure 6 Ordination diagram (DCA) of the ground beetle species assemblages of the Carpathians and ad-
jacent regions derived from 160 sample plots and 199 species. Axis 1: eigenvalue 0.64, length of
gradient 7.25. Axis 2: eigenvalue 0.38, length of gradient 4.33.
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Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

South Alps and West Balkans. Some species Discussion

occurred only in beech forests in the Carpathi-
an and Illyric regions, respectively (Fig. 5, Ta- Mesoscale diversity
ble 4). Carabid assemblages in beech forests
of the Carpathians and adjacent regions were Late Holocene mass expansion of beech and the
more distinct, showing very little overlap with spread of zonal beech forests to West Central
those of other habitat types (Fig. 6). Europa took place at the end of late Neolithic
We found 10 carabid species positively cor- (Lang 1994) and Bronze ages. Hence, interac-
related with beech cover in the Carpathians tions between beech and broadleaf forest spe-
and adjacent regions (Table 4). Some are Car- cies date back to just about 20 tree generations
pathian endemics (Carabus obsoletus subsp. (Walentowski & Winter 2013). The effective
carpathicus, Pterostichus foveolatus, P. pilo- time of coexistence may have been even short-
sus) or occur but very rarely in west central Eu- er due to simultaneous human impacts.
rope (Licinus hoffmannseggi). Further species Among the 48 beechwood dwelling PFR
(Abax parallelepipedus, Cychrus caraboides) species of saproxylic beetles and the beech
showed a positive correlation with beech forest ground-dwelling carabid and mollusc
abundance but are not beech forest specialists species observed in our study, neither exclu-
in West Central Europe. None of these species sive beech forest specialists nor regional en-
necessarily is tied to beech forests, let alone demics were found. This is an indicative of a
in all parts of its range. For example, even P. rather short period of joint evolution in beech
pilosus as a Carpathian endemic can be found ecosystems. Recent speciation, as observed in
in sub-alpine areas and high-elevation spruce the land snails Bulgarica vetusta subsp. festiva
forests, in our dataset it clearly favours beech and Cochlodina costata subsp. franconica on
forests in the way that it is positively corre- dolomite and limestone rocks and boulders
lated to the share of beech. Regionality of the (Hässlein 1960), seems to be restricted to more
habitat may play a major role, especially for or less open habitats. This result is congru-
those species whose range extends beyond one ent with findings in forest systems regarding
region, but possibly even within one region if vascular plants (Walentowski & Zehm 2010)
it is large enough in longitudinal or latitudinal and woody plants (endemic Sorbus apomicts,
extension and if subspecific taxa are involved. Meyer et al. 2005), and also with the ground

Table 4 Ground beetle (carabid) species positively correlated with beech dominance as derived from a
CCA of the macroscale dataset (threshold r > 0.4).
r - value t - value df p - value
Carabus auronitens 0.59 9.09 158 < 0.001
Carabus obsoletus 0.52 7.64 158 < 0.001
Molops piceus 0.50 7.19 158 < 0.001
Carabus linnei 0.49 6.99 158 < 0.001
Abax parallelepipedus 0.46 6.55 158 < 0.001
Cychrus caraboides 0.45 6.38 158 < 0.001
Pterostichus foveolatus 0.45 6.34 158 < 0.001
Pterostichus burmeisteri 0.44 6.13 158 < 0.001
Pterostichus pilosus 0.44 6.08 158 < 0.001
Licinus hoffmanseggi 0.40 5.45 158 < 0.001
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Ann. For. Res. 57(2): 215-231, 2014 Research article

beetles to be found in the canopy of European deadwood retreated to isolated locations or be-
beech forests (Müller-Kroehling & Zehetmair came extinct.
2014). Our data reveal that the majority of the
Specific habitat structures and habitat conti- ground-dwelling carabids and molluscs of the
nuity seems to be more important for the spe- beech forests in West Central Europe are in-
cies groups investigated than the presence or deed euryoecious, i.e. to be found in a variety
abundance of beech. Saproxylic PFR beetles of forest habitats. It can be assumed that they
have survived on deadwood of various host were already present prior to the immigration
tree genera, some confined to ancient trees in of beech in mid-Holocene mixed deciduous
pasture landscapes and parks (Speight 1989, broadleaf forests of Ulmus, Tilia, Fraxinus
Warren & Key 1991), partly substituting for and Quercus, as indicated by the large faunal
the loss of habitats in primeval forests. In similarities. Specifically, the more stenoecious
fact, temporal scale as well as structure and taxa among the ground-dwelling mollusc and
surroundings matter for the survival of relict ground beetle species require deadwood and
populations (Økland et al. 1996, Jonsson et al. stand continuity, which may be provided by
2005, Sverdrup-Thygeson et al. 2010, Buse shady deciduous forest types in general (e.g.,
2012, Sverdrup-Thygeson et al. 2014), and ravine forests), not exclusively by beech for-
many of the PFR xylobiont beetles in Europe ests. Fossil evidence suggests that woodland
are rather primeval forest structure specialists snail diversity declined significantly during
than primeval forest species. Even the ceram- the late Holocene (Hässlein 1960, Ložek 1982
bycid Rosalia alpina (EC Habitats Directive, a, b, c, Gedda 2006). Beech expansion might
Supporting Information II and IV), often cited have been a decisive factor for that, due to the
as flagship species of old beech forests, is ra- unfavourable chemical composition of leaf lit-
ther a specialist for sun-exposed trunks of vari- ter, the production of acid stemflows and the
ous broad-leaved trees (Fagus, Fraxinus, lowering of soil pH values and microbial acti-
Tilia, Ulmus, Acer) featuring dry decaying vity (Jacob et al. 2010, Thoms et al. 2010). The
timber in ancient woods and grazed open land snails Azeca goodallii and Perforatella
forests (Russo et al. 2010). bidentata were more common in the Atlan-
Many forest specialists are poor dispersers tic period from about 7500 to 5000/4500 cal
due to the relative stability of their habitat and BP, and under present-day conditions in Ba-
were therefore little likely to spread from their varia occur only in few residual populations
refuge areas to the establishing beech forests (Hässlein 1960). They are presently associated
of Central Europe. A high degree of overlap with moist boulder scree forest with Tilia, Ul-
in the invertebrate communities of different mus, Fraxinus and Acer, under conditions that
forest types in West Central Europe suggests do not allow beech to become dominant.
that many species of beech forests observed
today originate in other – adjacent – types of Macroscale diversity
woodland such as oak forests and in particu-
lar tree species-rich ravine and boulder forests. Pyrenees, Alps, Apennines and Carpathians are
Due to massive human impacts on woodlands hotspots of beech forest endemics and stenoe-
such as historic overexploitation, an overall al- cious species (Pott 2000, Willner et al. 2009).
teration and simplification of forest structures While these mountain ranges form a massive
caused by post-war depletion and acquisitive East-West barrier inhibiting species migrations
forestry, especially in the past 60 years, PFR in the Quaternary, they also provided refuges in
species that depend on habitat continuity and sheltered places such as in steep-sided valleys,
structural diversity including high amounts of and triggered speciation (Magri 2008). The

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Walentowski et al. Faunal diversity of Fagus sylvatica forests ...

Balkan and the Rhodope Mountains facilitated Italian montane forests have 159 ground bee-
species migrations between the biodiverse Il- tle species endemic to Italy, corresponding to
lyric and Carpathian beech forests and the an endemism rate of 31% (Brandmayr et al.
Mediterranean and Black Sea regions (Willner 2003).
et al. 2009, Walentowski et al. 2010). Short An increase in endemism towards the south-
distances to refuges allowed the expansion of ern and southeast European mountains is evi-
broadleaf forests including beech during inter- dent also for terrestrial molluscs (Cameron et
and postglacial periods. Coherent forest areas al. 2011). Hotspots of mollusc endemism are
with primeval beech forest remnants occurred found in the Mediterranean and Alpine biogeo-
throughout the Holocene. Intact predator-prey graphic regions, especially on limestone sub-
relationships ensured the processual integrity strates. These overall trends have been speci-
of forest. fied in a case study on forest ground-dwelling
Our results suggest a species-time or host- molluscs from the Slovak Carpathians (Čiliak
age dependent relationship (Southwood 1961, & Šteffek 2011). The authors observed among
Birks 1980, Rosenzweig 1995) between inver- others seven Carpathian endemic gastropods
tebrate assemblages and beech ecosystems, as (Bielzia coerulans, Cochlodina cerata, Faus-
shown above for saproxylic beetles in southern tina faustina, Monachoides vicinus, Perfor-
Romania. The patterns of richness and ende- atella dibothrion, Vitrea transsylvanica, Vestia
mism among the investigated beetle groups gulo), one Alps-Carpathian disjunction (Euco-
confirm the validity of the refuge distance bresia nivalis) and one land snail restricted to
model found for vascular plants (Willner et the Western Carpathians (Petasina unidentata
al. 2009). Ground beetle assemblages at short subsp. carpatica), suggesting a glacial forest
distance from potential refuge areas contain refuge that remains to be confirmed.
more species associated with beech forest (as-
certained by our data for 10 carabids in the
Carpathians) and narrow range species (three Conservation issues
Carpathian endemics). None of these species is
exclusive to beech forests, let alone in all parts Conservation status assessments based merely
of its range. For example, Pterostichus pilosus, on species numbers may fall short if the dis-
a Carpathian endemic (Nitzu et al. 2008), oc- tinctiveness of species pools goes unregarded
curs in sub-alpine areas and high-elevation (Andersson et al. 2013). Our study on Euro-
spruce forests, but in our dataset is positively pean beech forest highlights the importance of
correlated to the proportion of beech. Region- studying macroscale biodiversity to see mes-
ality of the habitat does play a major role, es- oscale biodiversity in perspective. On the one
pecially for those species whose range extends hand this perspective corroborates the neces-
beyond one region, but possibly even within sity of international conservation strategies,
one region if it is large enough in longitudinal on the other hand conservation responsibilities
or latitudinal extension and if subspecific taxa and priorities might differ at a regional level.
are involved. Macroscale responsibilities requiring na-
Studies such as of Brandmayr (1983), Co- tional and international conservation efforts
mandini & Vigna Taglianti (1990), Vigna Ta- for European beech forest should focus on the
glianti & De Felici (1994) and Rizun & Chu- preservation of sufficiently large and coherent
mak (2003) showed that the Carpathians, but areas in the core regions of European beech
also the Pyrenees, Southern Alps, Apennines with special attention to the south, south cen-
and Balkan ranges are home to a carabid fauna tral and southeast European mountains with
of high species diversity and endemic richness. glacial refuge areas. These eco-historically old

227
Ann. For. Res. 57(2): 215-231, 2014 Research article

beech forest areas and, in particular, the unique of climate change pressure on cold-preferen-
primeval forests, are essential for most of the tial species (Müller-Kroehling et al. 2014).
beech forest dwelling endemics and stenocious As PFR species occur in isolated sites and
species, thus exhibiting a maximum evolution- depend chiefly on wood amounts and tree
ary and ecological distinctiveness. structures and less on beech forest as such,
Mesoscale responsibilities requiring re- their preservation also relies on local efforts in
gional, national or binational efforts in ecohis- small areas, sometimes focussing on specific
torically young beech forest areas, e.g. in Ger- tree groups, sometimes on exposed wooded ar-
many, should, in the absence of narrow-range eas or on azonal formations. Ongoing manage-
endemics and with only very few beech spe- ment directed to keep old oaks free standing
cialists, focus on maintaining and improving and sun exposed and to ensure the recruitment
forests composed of native tree species in their of young oaks, and the restoration of aban-
natural habitats, by designing sizable forest doned pasture woodlands should have a high
reserves containing considerable proportions priority in nature conservation (Jansson 2009).
also of non-beech forest types. Also the beech forest natural ecotones require
In west central European beech forest eco- special attention as they may provide habitats
systems, being by far the most important for particular relict woodland invertebrate
broadleaf forest type, their local conditions, populations.
processes and functions have ever been much
affected by human land use and forestry.
Conservation focus should be on the faunal, Acknowledgements
fungal and floral diversity of old-growth for-
ests in general. Saproxylic beetles, woodland We thank Volodymyr Rizun, Department
ground beetles and woodland molluscs of the of Regional Biocoenotic Monitoring of the
young post-ice age temperate forests, that are National Academy of Sciences of Ukraine,
presently labelled as beech forest species are Wolfgang Waitzbauer (University of Vienna),
in fact linked to an adequate frequency and Ferenc Kádár, and Heiko Sparmberg for cor-
habitat continuity of admixed secondary tree respondence and contributions to the ground
species (like Quercus, Acer, Fraxinus, Ulmus) beetle data. We further thank anonymous re-
within beech forest landscapes. For cool tem- viewers for competent advice on previous ver-
perate mountain mixed forests with distinct sions of the article.
boreal-montane species pools (e.g. related to
Urwald relict species) and species which are
old growth obligates (like the ground beetle References
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