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Influence of tree plantations on the

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DOI: 10.1016/j.foreco.2016.06.011

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 Forest Ecology and Management 376 (2016) 231237

Contents lists available at ScienceDirect

Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Influence of tree plantations on the phylogenetic structure

of understorey plant communities
Marcin Piwczynski , Radosaw Puchaka, Werner Ulrich
, Poland
Chair of Ecology and Biogeography, Nicolaus Copernicus University, Lwowska 1, PL-87-100 Torun

a r t i c l e i n f o a b s t r a c t

Article history: The effect of non-native and native tree species plantations on plant understorey diversity and function-
Received 9 February 2016 ing has gained major interest among ecologist and conservationists. The majority of studies, however, has
Received in revised form 3 June 2016 been focused on changes in species richness and composition in plantations in comparison with natural
Accepted 6 June 2016
forests. Recently, the phylogenetic community structure (PCS), i.e. the degree of phylogenetic relatedness
within a community, came into the focus of interest. As species richness and PCS are not equivalent, the
question of how plantations change PCS and thus the evolutionary history conserved in the associated
Keywords:
understorey community is important to fully assess the biodiversity of tree species introductions. In this
Net relatedness index
Nearest taxon index
study we used phytosociological data from the Cedynia Landscape Park in western Poland containing
Species co-occurrence anthropogenic mono-cultures of Scots pine and introduced trees (black locust and Douglas fir) as well
Ellenberg values as natural sessile oak stands to study the influence of plantations on species richness, composition,
Natural oak forest and phylogenetic diversity of understorey vegetation. We also added relevs from mixed forests naturally
occurring on wetter and richer soil to obtain the full regional species pool. Our results revealed that plan-
tations do not consistently change PCS; they either increased (Douglas fir) or decreased (black locust) the
phylogenetic relatedness of understory plant communities. Moreover, our data suggest that environmen-
tal conditions are more important drivers of understorey community composition than dominant tree
identity. In this respect we identified light regime, pH, and nitrogen availability as major drivers of phy-
logenetic diversity.
2016 Elsevier B.V. All rights reserved.

1. Introduction functioning has gained interest among ecologist and conservation-

Anthropogenic changes in plant community structure, for
instance the plantation of non-native trees, can disturb the native
flora (Fisher and Owens, 2004; Kotiaho et al., 2005) and operate as
an additional environmental filter that affects understorey species
composition (e.g. Williams et al., 2009; Sullivan et al., 2015). Both
these factors might cause changes in the distribution of plant char-
acters (Ding et al., 2012; Feng et al., 2014; Larkin et al., 2015) and
community functioning (Lavorel and Garnier, 2002; Mouillot et al.,
2013). As understories play a central role in the dynamics and
functioning of forest ecosystems by contributing to forest nutrient
cycles (Chastain et al., 2006; Zhao et al., 2013), productivity (Chen
et al., 2004; Reich et al., 2012), and regeneration ability (Nilsson
et al., 2005; Royo and Carson, 2006), the potential negative effect
of non-native tree species on plant understorey diversity and
Corresponding author.
E-mail addresses: piwczyn@umk.pl (M. Piwczynski), puchalka@umk.pl
(R. Puchaka), ulrichw@umk.pl (W. Ulrich).
ists (e.g. Bremer and Farley, 2010). Although plantations are some-
times seen as green deserts in comparison with natural forests
(Bremer and Farley, 2010) recent work highlighted the potential
positive effect as a catalyst of vegetation recovery (Butler et al.,
2008; Barbosa et al., 2009; Mendona-Lima et al., 2014).
Environmental stress might induce changes of another form of
diversity, the phylogenetic community structure (PCS), i.e. the
degree of phylogenetic relatedness within a community (Webb
et al., 2002). Ecological experiments have shown that communities
made of evenly or distantly related species were more stable and
productive than those with more clustered phylogenetic relation-
ships (Cadotte, 2013; Cadotte et al., 2012). Additionally, phyloge-
netically diverse communities were shown to support more
species at various trophic levels (Dinnage et al., 2012). As species
richness and PCS are not equivalent (Devictor et al., 2010; Pio
et al., 2011) the question of how non-native tree introduction
changes PCS and thus the evolutionary history conserved in the
associated understorey community is important to fully assess
the subsequent effects on forest biodiversity and functionality
(Cadotte et al., 2012; Cadotte, 2013).

http://dx.doi.org/10.1016/j.foreco.2016.06.011
0378-1127/ 2016 Elsevier B.V. All rights reserved.
232 ski et al. / Forest Ecology and Management 376 (2016) 231237
M. Piwczyn

Differences in PCS between natural and planted stands alone identical potential natural vegetation (Pawlaczyk and Kujawa-
are not easily traced back to the process of species assembly due Pawlaczyk, 1999; Puchaka and Pachocki, 2014). Focusing on dif-
to the interplay of species interactions and abiotic forces. Analyses ferences in PCS these unique conditions allowed us to ask whether
of species interactions have traditionally focused on competition as (1) plantations of non-native and native trees introduce strong
the major force shaping patterns of species co-occurrences (Grime, habitat filters which decrease phylogenetic relatedness of under-
1973; Diamond, 1975). Darwins (1859) argument that closely storey communities in comparison with natural tree stands,
related species compete stronger than their distantly related coun- whether (2) the possible changes in PCS correlates with respective
terparts (the competition-relatedness hypothesis) has become a changes in species richness and composition, and (3) how forest
central tenet of community ecology (Svenning et al., 2008; Allan specific soil factors influence PCS and species richness.
et al., 2013), although Cahill et al. (2008) found little direct evi-
dence for this assertion. In connection with the competitive exclu-
2. Methods
sion principle Darwins hypothesis predicts that natural
communities should contain more evolutionary lineages than
2.1. Data collection and reconstruction of phylogeny
expected from a random assembly taken from the regional species
pool (phylogenetic overdispersion, Webb et al., 2002). In turn, abi-
We collected phytosociological data from the Cedynia Land-
otic factors are often considered as filters that allow closely related
scape Park (Pawlaczyk and Kujawa-Pawlaczyk, 1999; Pakalski
species to coexist because they are more likely to share similar
et al., 2005) to construct a presence/absence table containing a
traits (reviewed in Mayfield and Levine, 2010 and Gtzenberger
total of 171 understorey plant species in rows and 96 sites in col-
et al., 2012) thereby linking the concepts of niche conservatism
umns (electronic supplement Table A1). All sites were located
(the sharing of traits by closely related species, Wiens et al.,
within sandy areas (acid and bleached brown soils) naturally char-
2010) with phylogenetic clustering (a lower phylogenetic diversity
acterized by oak forest (Calamagrostio arundinaceae-Quercetum;
than expected from a random sample, Webb et al., 2002). There-
Pawlaczyk and Kujawa-Pawlaczyk, 1999, Fig. 1) and represented
fore, environmental filtering should reduce the number of lineages
one phytosociological relev of 100400 m2 area.
and consequently phylogenetic relatedness (Wiens et al., 2010).
A total of 32 sites occurred in (semi)natural oak forests (Quercus
Recent studies have shown that biotic factors such as competitive
petraea (Matt.) Liebl.), 21 in native Scots pine (Pinus sylvestris L.), 3
interactions between distantly related species may also promote
in Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) and 10 in black
the coexistence of closely related species (Mayfield and Levine,
locust (Robinia pseudoacacia L.) plantations. In order to cover the
2010).
total pool of regionally occurring understorey species we added
In our study, we applied phylogenetic diversity metrics to esti-
30 relevs from mixed forests naturally occurring on wetter and
mate the impact of tree plantations on the structure of understorey
richer brown soils. To assess habitat conditions we used five Ellen-
plant communities. This impact can be manifold; each particular
berg indicator values: light, temperature, soil moisture, soil reac-
tree species can alter a number of abiotic factors such as light
tion (pH), and soil nitrogen (Ellenberg et al., 1992) as provided
regimes, soil water availability, soil nutrients, or the physical prop-
by JUICE 7.0 (Tichy, 2002). Average Ellenberg values per site were
erties of litter (Augusto et al., 2002, 2003; Barbier et al., 2008). The
calculated over all species present at a site.
replacement of native forests by tree plantations introduces new
We established the phylogenetic relationships among all under-
environmental filters and simultaneously removes others. This
storey species by using the Phylomatic software, a phylogenetic
has an effect on species composition and richness. Many authors
database and toolkit for the assembly of phylogenetic trees
have found a decrease in species richness in plantations in compar-
(Webb and Donoghue, 2005). Trees generated by the software
ison with (semi)natural communities (e.g. Van Wesenbeeck et al.,
were based on APG III (Angiosperm Phylogeny Group, 2009). We
2003; Goldman et al., 2008; Paritsis and Aizen, 2008; Bremer and
used peer reviewed molecular phylogenies to resolve most of the
Farley, 2010; Proena et al., 2010; Quine and Humphrey, 2010)
remaining polytomies, leaving mainly those inside genera (for a list
while cases of increase or equivalence seem to be uncommon
of publications and the tree in Newick format see Table B1). We
(Bremer and Farley, 2010; Quine and Humphrey, 2010; Li et al.,
assigned branch lengths to the tree using the BLADJ (Branch Length
2014). Exotic understorey plants with broad niche breadth are
Adjustment) option in Phylocom (Webb et al., 2008). Branch length
often much more abundant and diverse in plantations than in nat-
was determined by assigning a minimum age to genera, families,
ural forests (Bremer and Farley, 2010). Additionally plantations fre-
and higher orders from the molecular dating of Wikstrm et al.
quently have a reduced diversity in indigenous plants (e.g.
(2001) and spacing undated nodes evenly between dated ones.
Goldman et al., 2008), although the spatial proximity to natural
Since Wikstrms dating does not include ferns, we used ages gen-
forests and the age of the managed plantations can strongly influ-
erated by Schuettpelz and Pryer (2009) to assign them to nodes in
ence this trend due to altered seedling recruitment (Lee et al.,
the fern part of the phylogeny.
2005; Farwig et al., 2009). Because alien tree species modify exist-
ing environmental filters they should also influence species rich-
ness and composition, and consequently PCS. However, 2.2. Comparison of the natural oak forest and tree plantations
respective comparisons have, to the best of our knowledge, not
yet been attempted. To infer the differences in PCS, we used two approaches. First,
Particularly little is known about the influence of soil conditions we calculated the net relatedness index NRI, which is the standard-
on the PCS of plants. Schreeg et al. (2010) found that phylogenetic ized mean phylogenetic distance (MPD) (Webb et al., 2002). NRI
structure in a tropical tree community varied along gradients of relies on a particular null assumption to which raw scores are com-
soil conditions and water availability and Ulrich et al. (2014) pared. In its original definition, it is the negative standardized
reported low soil carbonate content to drive communities to phy- effect sizes (SES) of MPD with respect to the predefined null model
logenetic overdispersion. Therefore it is important to account for distribution (Webb et al., 2002). Under the assumption of a normal
soil variables in comparative studies of introduced and native tree error structure SES scores should have values below
1.96 and
stands. In this study we use phytosociological data from the Cedy- above +1.96 at the two-sided 5% error level. For comparability with
nia Landscape Park in western Poland (Fig. 1) containing anthro- other diversity metrics we here use the complement -NRI that
pogenic monocultures of introduced trees as well as natural oak increases with increasing phylogenetic diversity. Second, we esti-
stands. All stands share a similar sandy soil type and have an mated the effect of introduced and native tree plantations on the
 ski et al. / Forest Ecology and Management 376 (2016) 231237
M. Piwczyn 233

Fig. 1. Location of the study area in Poland (A). Distribution of forest sites within the Cedynia Landscape Park (53050 N53450 , 14070 14380 E, 839.70 km2) (B) natural oak
forest, (C) scots pine plantations, (D) black locust plantations, (E) Douglas fir plantations and (F) mixed forest. Grey denotes areas for which oak forest is the potential natural
vegetation (Pawlaczyk and Kujawa-Pawlaczyk, 1999).

phylogenetic structure of understorey plant communities using the
nearest taxon index NTI (Webb et al., 2002). NTI quantifies the
phylogenetic relatedness of the endpoints of the phylogenetic tree.
Again we use -NTI for comparability. NRI and NTI are partly com-
plementary as NTI is focused on the terminal part of the phyloge-
netic tree and fine-scale relatedness, while NRI covers more basal
parts and deeperlevel diversity (Webb et al., 2002). Indeed when
applied to our data NRI and NTI were only moderately positively
correlated (r2 = 0.37).
At higher species richness, the probability of including small
and distantly related taxa in the community and consequently
MPD increases. To eliminate this bias in statistical inference appro-
priate null or neutral models have to be applied (Gotelli and Ulrich,
2012). In the present case, we compare the observed species com-
position with a random sample from same source pool. As there is
a priori no constraint on local species richness a suitable null
hypothesis is the random appearance of species in single sites
constrained only by differences in regional species abundances
and therefore colonization probability. This constraint was
approximated by the observed total number of occurrences in all
sites in a given year. Therefore we used a null model that resam-
ples the species  sites matrix proportional to observed species
occurrences and total species richness per site (the marginal occur-
rences totals) as recommended by Ulrich and Gotelli (2012).
We used non-metric multidimensional scaling (NMDS) based
on Soerensen dissimilarity among the 96 stands (as implemented
in Canoco 5) for inferring differences in species and phylogenetic
composition between the forest types (monocultures versus mixed
stands, natural versus plantation). We compared species richness,
forest type, and PD using one- and two-way Permanova
(Anderson, 2001) based on Euclidean distances as implemented
in Primer 7. Finally we used general linear modelling (orthogonal
sums of squares) to relate species richness and PD to forest type
and habitat characteristics (Ellenberg values) across stands. Post
hoc comparison of categorical variables were based on unequal
sample size Tukey tests. To exclude possible effects of area on
relev species richness but also the possible richness effect on
the phylogenetic metrics our regression analyses always included
234 ski et al. / Forest Ecology and Management 376 (2016) 231237
M. Piwczyn

richness as a covariate. All significance levels were Bonferroni 0.2

corrected.
0.15
3. Results 0.1
T
0.05 M
One-way Permanova detected highly significant differences

NMDS 2
L N
between all forest types with regard to Ellenberg habitat condi- 0
tions (pseudo F = 32.3, P < 0.0001). Post hoc pairwise comparisons -0.2 -0.1 0 0.1 0.2
revealed for all pairs significant habitat differences (P < 0.05, not -0.05 pH
shown). The mixed forest stands had the highest moisture, pH, -0.1
and nitrogen Ellenberg scores (Table 1).
Forest types significantly differed in species richness (ANOVA -0.15
P < 0.0001) although type explained less than 15% of variance in
-0.2
richness (Table 2). The pine and the mixed forests were most spe-
cies rich (Table 1) and significantly more diverse than the natural NMDS 1
oak and Douglas fir forests (Table 1, P < 0.01). NMDS analysis
Fig. 2. Non-metric multidimensional scaling separates species composition of the
(Fig. 2) separated the community composition of the mixed forest mixed forest stands (black circles) from the natural oak stands (red squares), black
from the four other forest types while among the mono-stands Locust (blue diamonds), Scots pine (green triangles), and Douglas fir (open circles)
only the natural oak forest and the Scots pine stands were signifi- by pH, nitrogen (N), moisture (M), temperature (T), and light (L) (stress = 0.22). The
cantly separated (Fig. 2; randomization P < 0.01). ellipses for each forest type include 95% of data points.
GLM detected significant differences in NRI and NTI between
the stands (P < 0.01, Table 2). Subsequent pairwise comparisons
Table 2
identified the black locust as being significantly (P < 0.05) phyloge- General linear mixed modelling identified forest type as to influence species richness,
netically clustered in comparison to the natural oak forest (Table 3). NRI (net relatedness index), and NTI (nearest taxon index). Given are variance
Black locust and Scots pine stands differed significantly in NRI partitioned partial coefficients of determination g2 and the respective coefficient of
(P < 0.01, not shown). After correction for sample area and species the total model.

richness (Table 2) forest type still had a significant influence on NRI Variable Effect df Species richness NRI NTI
and NTI (P < 0.01) although it explained less than 20% of variance Forest type Fixed 4 0.23*** 0.16** 0.18**
in each metric. None of the pair-wise post hoc comparisons of Sample area Random 4 0.14* 0.07 0.09
NRI and NTI (except of the Douglas fir black locust pair) remained Species richness Fixed 0/1 0.01 0.01
significant at P < 0.05 (not shown). Error 87/86

As forest type was linked to habitat characteristics (Table 1) we r2 (total model) 0.51*** 0.28** 0.27**
also compared PCS with respect to Ellenberg scores (Table 4). Spe- Bonferroni corrected parametric significances:
cies richness was positively linked to pH although pH explained *
P < 0.05.
**
only 6% of variance in richness. NRI was significantly negatively P < 0.01.
***
correlated with light and pH (P < 0.01) and positively with nitrogen P < 0.001.

(P < 0.05) demands while NTI was strongly linked to light (Table 4).
Table 3
4. Discussion NRI (net relatedness index) and NTI (nearest taxon index) scores ( one standard
error) of the six forest types and Bonferroni corrected significance levels of post hoc
Permanova pairwise comparisons with the natural oak forest.
Recent work on the influence of natural and human-induced
changes to communities have pointed to environmental filters as Site NRI NTI p(NRI) p(NTI)
the main factors affecting species assembly (Helmus et al., 2007; Natural oak forest 0.29 0.20 0.20 0.20
Dinnage, 2009; Pausas and Verd, 2008; Turley and Brudvig, Scots pine plantation 0.53 0.23
0.25 0.18 0.99 0.12
2016). The common conclusion from these studies is that such dis- Douglas fir plantation 1.75 0.76 0.93 0.60 0.32 0.29
Black locust plantation
0.91 0.38
1.33 0.36 0.06 <0.01
turbed communities on average contain more closely related spe-
Mixed forest
0.30 0.16 0.31 0.12 0.25 0.71
cies in comparison with undisturbed ones, because close relatives
have a higher probability of sharing similar functional traits. In
our study, however, the impact of plantations on the undergrowth respectively, although only the difference in NTI for black locust
revealed a variety of phylogenetic effects. Clearly, plantations do remained statistically significant after Bonferroni correction
not unequivocally reduce phylogenetic diversity. The effect on (Table 3). In the case of Douglas fir, the observed phylogenetic
total understorey richness depends on tree species identity overdispersion is apparently caused by a similar species composi-
(Table 1). tion across sampled sites: one or two fern species, one or two grass
Two alien species, Douglas fir and black locust showed higher species and Oxalis acetosella. All of these plants are distantly related
and lower phylogenetic relatedness than a natural oak forest, and induced high mean values of NRI and NTI (Table 3).

Table 1
Basic information about the forest types studied. Errors refer to standard errors.

Site Relevs Total area Species richness Ellenberg scores

Total Site Light Temperature Moisture pH Nitrogen
Natural oak forest 32 8600 33 5.41 0.44 5.44 0.60 5.15 0.06 4.97 0.03 3.29 0.03 3.89 0.11
Scots pine plantation 21 7025 69 10.05 1.20 6.28 0.10 5.31 0.07 4.85 0.06 4.13 0.22 4.87 0.26
Douglas fir plantation 3 675 5 3.33 0.72 4.21 0.31 5.00 0.00 5.00 0.00 3.36 0.16 4.39 0.16
Black locust plantation 10 3250 37 9.30 0.68 6.83 0.14 5.47 0.09 4.72 0.17 5.87 0.36 5.60 0.21
Mixed forest 30 10,100 115 15.10 1.16 4.71 0.09 5.15 0.14 5.35 0.09 6.06 0.17 5.81 0.17
 ski et al. / Forest Ecology and Management 376 (2016) 231237
M. Piwczyn 235

Table 4 2007; Devictor et al., 2010; Pio et al., 2011). As a general rule spe-
Multiple regression identified light and temperature as to influence species richness cies richness is thought to be an inaccurate surrogate of phyloge-
and soil pH and nitrogen as to influence NRI (net relatedness index) and NTI (nearest
taxon index). Given are partial g2 and b coefficients of determination and the
netic diversity. In our case, the pine plantations supported on
respective coefficient of determination of the total model. average more light and nitrogen demanding species from open
communities (Table 1) than the natural oak stands. For example,
Variable Species richness NRI NTI
the average Ellenberg values for nitrogen and light for 42 species
partial g2 b partial g2 b partial g2 b unique to the pine plantations and absent from the oak stands
Intercept 0.01 0.15 0.13** were 5.42 and 6.88, respectively, while 20 species unique to the
Species 0.01
0.10 <0.01
0.09 oak stands had values of 4.05 and 5.25, respectively. This finding
richness
implies increased species diversity after human disturbance
Light 0.05
0.21 0.12**
0.32 0.23***
0.50
Temperature 0.03 0.16 0.08
0.30 0.03
0.21 (Nagaike, 2002). Particularly forest management that invokes
Moisture <0.01
0.05 <0.01 0.05 <0.01 0.06 increased light regimes, for instance thinning, might effectively
pH 0.06* 0.38 0.21***
0.87 0.05
0.43 set back plant communities to an early successional stage with
Nitrogen 0.01 0.19 0.09* 0.51 0.06 0.42
many light demanding ruderal and invasive species (Swindel
r2 (total 0.41*** 0.34*** 0.25*** et al., 1986). The oak stands of our study were not managed in
model)
recent times while at least some pine plantations were regularly
Bonferroni corrected parametric significances: thinned. This difference might at least partly explain the differ-
*
P < 0.05. ences in species richness.
**
P < 0.01. Ratsirarson et al. (2002) and Stephens and Wagner (2007) have
***
P < 0.001.
argued that the introduction of native tree species should change
community composition of animals and plants to a lesser extent
than exotic ones, because native species are possibly a better
Interestingly, this higher phylogenetic variability compared to the equivalent of natural forest communities. Our study does not cor-
oak forest is probably caused by the stronger light regime in Dou- roborate this hypothesis. Understorey richness and PD of the two
glas fir plantations (Table 1) that filtered for plants with low light exotic plantations of black locust and Douglas fir did not consis-
demand (e.g. Pteridopsida, Schneider et al., 2004; O. acetosella, tently deviate from the natural Scots pine plantations (Tables 1
Packham and Willis, 1977). Apparently, the effective species pool and 3). Similar results have been obtained by Quine and
contains only a limited number of species with such low light Humphrey (2010) for exotic and native tree plantations in Britain,
demands, each of them evolved independently in unrelated lin- who argued that well-established exotic trees can even provide
eages. In spite of the higher shade tolerance of these species, they new habitat for otherwise endangered indigenous plant and fungal
are also able to co-occur in oak forests together with other, more species. We add that the local soil and light conditions seem to be
demanding species. more important for community assembly than a possible phyloge-
Unlike Douglas fir plantations, black locust plantations showed netic prefiltering by dominant tree species identity. Even at the
lower mean NRI and NTI scores than the natural oak stands indicat- mixed stands, in which species richness and composition differed
ing phylogenetic clustering (Table 3). Many species from the natu- considerably in comparison to the oak forests and plantations
ral oak forest adapted to low pH and nitrogen did not inhabit the (Table 1, Fig. 2), PCS did not significantly change.
Robinia forest. For example, there were no members of Asteraceae NTI and NRI cover partly contrasting patterns of phylogenetic
such as Hieracium, Solidago, Mycelis and the forest lacked some spe- diversity. Particularly we expected to see NTI, focusing on the
cies of monocots such as Convalaria majalis, Maianthemum bifolium, fine-scale structure of community composition, to react on light
Polygonatum odoratum, Luzula pilosa, which are indigenous for nat- regime and soil type. This was only partly the case (Tables 2 and
ural oak and pine forests in this region (Ellenberg, 2009; 4). While both metric identified the differences in PCS between for-
Matuszkiewicz, 2011). Our finding thus corroborates the observa- est types in a similar manner, both differed with respect to envi-
tion of a decreasing number of species naturally occurring in for- ronmental conditions (Table 4). We interpret these results in a
ests in favour of exotic and ubiquitous ones in plantations way that deeper rooted patterns of PCS are linked to soil properties
(Bremer and Farley, 2010). (pH, nitrogen), while co-existence of closely related species is gov-
The black locust plantation was dominated by grass species, erned by light regimes (Table 4) with shadow sites favoring phylo-
which comprise over 40% of all species (versus 19% in oak forest, genetic overdispersion.
Table A1). There were over 50% of grass species in 6 from 10 sites.
Likely, a simultaneous decrease of the light regime and an increase
of soil nitrogen (Table 1) due to N fixation of black locust allow 4.1. Conclusions
plants from open habitats with higher nitrogen and pH demands
(e.g. various species of grasses, Chaerophyllum temulum, Chelido- Our study demonstrated mixed influences of exotic and native
nium majus, Anthriscus caucalis) to colonize black locust plantations tree plantations on species richness, composition and phylogenetic
(Vtkov and Kolbek, 2010). diversity of the associated understorey vegetation. Plantations and
The phylogenetic diversity of the native Scots pine plantations native oak stands were not strongly separated with respect to spe-
was not significantly different from the natural oak forest and also cies richness and phylogenetic structure. Our data suggest that
from the randomly assembled null model communities (Table 3). local environmental conditions are more important drivers of
On the other hand, we found significant differences in species understorey community composition than dominant tree identity.
number (Table 1) and species composition (Fig. 2). This shows that In this respect we identified light, pH, and nitrogen availability as
species rich communities do not have automatically equally high major drivers of phylogenetic diversity.
levels of phylogenetic diversity since species share evolutionary
history with related species (which can be numerous or few, dis- Acknowledgments
tant, or close) (Devictor et al., 2010; Pio et al., 2011). Empirical
studies have shown that in some cases, patterns of species richness We would like to thank Maciej Pabijan and Hazel Pearson for
and phylogenetic diversity are closely correlated (Rodrigues and kindly improving our English. This research is linked to activities
Gaston, 2002), whereas in others they differ greatly (Forest et al., conducted within the COST FP1403 NNEXT network.
236 ski et al. / Forest Ecology and Management 376 (2016) 231237
M. Piwczyn

Funding: This work was supported by grants from the National Forest, F. et al., 2007. Preserving the evolutionary potential of floras in biodiversity
hotspots. Nature 445, 757760.
Science Centre, Poland, no. 2014/13/B/NZ8/04681 to WU.
Goldman, R.L., Goldstein, L.P., Daily, G.C., 2008. Assessing the conservation value of
a human-dominated island landscape: plant diversity in Hawaii. Biodiver.
Conserv. 17, 17651781.
Appendix A. Supplementary material Gotelli, N.J., Ulrich, W., 2012. Statistical challenges in null model analysis. Oikos
121, 171180.
Gtzenberger, L. et al., 2012. Ecological assembly rules in plant communities
Supplementary data associated with this article can be found, in
approaches, patterns and prospects. Biol. Rev. 87, 111127.
the online version, at http://dx.doi.org/10.1016/j.foreco.2016.06. Grime, J.P., 1973. Competitive exclusion in herbaceous vegetation. Nature 242, 344
011. 347.
Helmus, M.R., Bland, T., Williams, C., Ives, A., 2007. Phylogenetic measures of
biodiversity. Am. Nat. 169, 6883.
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