Science of the Total Environment 408 (2010) 5506–5512
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Science of the Total Environment
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / s c i t o t e n v
Oak forest exploitation and black-locust invasion caused severe shifts in epiphytic
lichen communities in Northern Italy
Juri Nascimbene a,⁎,1, Lorenzo Marini b,1
a
b
Department of Biology, University of Padova, via U. Bassi 58/B, I-35121 Padova, Italy
Department of Environmental Agronomy and Crop Production, University of Padova, viale dell'Università 16—35020 Legnaro, Padova, Italy
a r t i c l e
i n f o
Article history:
Received 30 April 2010
Received in revised form 17 July 2010
Accepted 19 July 2010
Available online 14 August 2010
Keywords:
Alien species
Cyanolichens
Forest management
Rare species
Trentepohlioid lichens
a b s t r a c t
In the last two centuries, native European oak forests have undergone a dramatic decline related to
increasing human pressure for agriculture and urbanization. Oak forests were either completely eradicated
and transformed into agricultural landscapes or replaced by second-growth formations. Intensive forest
management and the replacement of native forests with production forests or arable lands are recognized
amongst the main threats to many lichens in Europe. In this study, we used historical information on the
epiphytic lichen biota which was hosted in a native oak-dominated forest of Northern Italy to identify shifts
of lichen communities due to the changes in land use which occurred during the last two centuries. We also
compared the epiphytic lichen communities inhabiting remnant oak forests with those found in the habitats
that have replaced native forests: black-locust forests and agrarian landscapes. Almost all the species
sampled during the 19th century are now extinct. The loss of native habitat and the subsequent invasion by
black locust were probably the most influential factors which affected the composition of lichen
communities, causing the local extinction of most of the species historically recorded. Despite the fact
that oak remnants host only a few species which were historically recorded, and that they currently are the
lichen poorest habitat in the study region, they host lichen assemblages differing from those of black-locust
forests and agrarian stands. In these habitats lichen assemblages are mainly composed of species adapted to
well-lit, dry conditions and tolerating air pollution and eutrophication. This pattern is likely to be common
also in other lowland and hilly regions throughout Northern Italy where oak forests are targeted among the
habitats of conservation concern at the European level. For this reason, a national strategy for biodiversity
conservation and monitoring of lowlands forests should provide the framework for local restoration projects.
© 2010 Elsevier B.V. All rights reserved.
1. Introduction
The exploitation of forests may produce severe impacts on several
components of biodiversity (Heywood, 1995). In the last two
centuries, native oak forests across whole Europe have undergone a
dramatic decline related to increasing human pressure for agriculture
and urbanization. Native oak forests were either completely eradicated and transformed into agricultural landscapes or replaced by
second-growth formations. Black locust (Robinia pseudoacacia L.), an
alien species introduced in the late 1700s from the southeastern
United States, is amongst the most invasive species that have replaced
oaks in Italy (see e.g. Caprio et al., 2009; Motta et al., 2009). The high
potential for wood and erosion control, together with its fast and
highly competitive growth, greatly enhanced the spread of this
species which invaded large wet and fertile areas.
Amongst the various components of forest biodiversity, epiphytic
lichens are considered particularly sensitive to human-related altera⁎ Corresponding author. Tel.: +39 43942894.
E-mail address: junasc@libero.it (J. Nascimbene).
1
Both authors equally contributed to the study.
0048-9697/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.scitotenv.2010.07.056
tions of forests (Aragón et al., 2010) such as intensive management or
replacement of native forests with secondary forests (e.g. Fritz et al.,
2008; Hedenås and Ericson, 2000; Humphrey et al., 2002; Johansson,
2008; McCune, 2000; Moning and Müller, 2009; Nascimbene et al.,
2007; Rogers and Ryel, 2008). For instance, clearing of forests and
logging can alter humidity, temperature and light conditions (Moen
and Jonsson, 2003), causing a systematic reduction in populations and
local extinction of some cyanolichens and some crustose sorediate
species (e.g. Aragón et al., 2010). Management also tends to diminish
some types of phorophyte in the landscape, and consequently, the
substrate specialist lichens also decrease (Hedenås and Ericson, 2008;
Belinchón et al., 2009). Moreover, forest management and land-use
changes might interplay with the negative effects of air pollution (e.g.
Geiser and Neitlich, 2007; Jeran et al., 2007; Jovan and Mc Cune, 2004;
Mayer et al., 2009) and climate changes (e.g. Ellis and Coppins, 2007;
Ellis et al., 2007, 2009) resulting in a severe decline of several sensitive
species such as the two large macrolichens Usnea longissima (e.g.
Esseen et al., 1981; Nascimbene and Tretiach, 2009) and Lobaria
pulmonaria (e.g. Scheidegger and Werth, 2009).
Science-based assessment of forest lichen decline is often hindered
by the scarcity of detailed historical information available for direct
J. Nascimbene, L. Marini / Science of the Total Environment 408 (2010) 5506–5512
comparison with the communities found in the remnant habitats.
Historical comparisons have been successfully applied only within
urban environments, where site-referred historical data could be
more easily retrieved (Rose and Hawksworth, 1981; Isocrono et al.,
2007). When this information is available, historical comparison is a
powerful tool to evaluate changes in biodiversity (Biesmeijer et al.,
2006; Mac Nally et al., 2009), despite some limitations, related to
heterogeneity of available datasets or differences in the sampling
effort (e.g. Kosior et al., 2008; Newbold, 2010). In this study, we used
historical information on the epiphytic lichen biota which was hosted
in native oak forests in a region in Northern Italy. These data provide
an exceptional framework to identify shifts in lichen communities due
to the changes in land use which occurred during the last two
centuries.
Specifically this study had two main objectives: (1) to check if
epiphytic lichens historically recorded (1851–1894) in a region
formerly covered by large native oak forests are still present in the
remnant forests, and (2) to compare the epiphytic lichen communities inhabiting the remnants, with those found in the habitats that
have replaced the native forests (black-locust forests and isolated
trees in agrarian landscapes).
2. Materials and methods
2.1. Study area
The study was carried out in the Montello area, an elliptical shaped
hill placed in the northern part of the Venetian plain (NE-Italy, Veneto,
Treviso), extending over a 5000 ha surface with its main axis (13 km)
NNE–SSW oriented (Fig. 1). The mean elevation is 200 m (maximum
369 m) and it is 100 m higher than the surrounding plain (Ferrarese
and Sauro, 2005). The climate is temperate to humid subtropical with
a mean annual temperature of 12.9 °C, January being the coldest
month (mean temperature 3–4 °C) and July being the warmest one
(mean temperature 22.7 °C). Mean annual rainfall is 1100 mm, with
maxima in spring and autumn.
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2.2. Historical landscape and forest management
For centuries the Montello area was covered by a continuous oak
[Quercus robur L. and Q. petraea (Matt.) Liebl.] forest. However,
similarly to what occurred over much of the hilly and lowland
deciduous forests in Northern Italy, since the end of the 19th and
during the 20th centuries native oak forests were subjected to severe
cutting for agriculture land reclamation.
The first written information on the forest of Montello is available
in a document dated 994 (Favero, 1875). From 1389 to 1797 the forest
was owned by Venice republic and was used for providing the wood
for the huge naval arsenal. The management was based on selective
cutting resulting in a continuous, pure multilayered oak forest
(Favero, 1875; Susmel, 1994). After the decline of Venice the forest
was owned by the Austro-Hungarians until 1866. In this period, the
intensity of the exploitation increased mainly due to illegal cutting by
the local population. The intense disturbance promoted also the
establishment of black locust (R. pseudoacacia), which was favoured
by man because of its fast growth and good wood quality for firewood
and agricultural uses. In 1892, the local stakeholders declared the total
destruction of the native forest and the conversion of the whole area
into a farmland (Bertolini, 1905). However, due to non-optimal
conditions for agriculture, the conversion was not completed and part
of the area remained covered by forests which were overexploited
and greatly disturbed during the First World War. Between 1950 and
1970 the area was largely abandoned while in the surrounding region
both industries and agriculture greatly developed. From 1954 to 2004
the total forested area progressively increased (Ferrarese and
Castiglioni, 2005), being mainly related to the rapid spread of black
locust, which prevented the re-establishment of the native oak forest.
2.3. Current landscape and management
Currently, Montello appears like an island surrounded by the
anthropogenic landscape of the Venetian plain with diffuse agriculture
and secondarily urbanization. In this area there are only minor industrial
Fig. 1. Geographic position of the study area in north-eastern Italy. In the detailed map, the position of the 18 sampling plots surveyed for comparing currently available habitats (oak
forest, agrarian habitat, and black-locust forest) for epiphytic lichens is reported.
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J. Nascimbene, L. Marini / Science of the Total Environment 408 (2010) 5506–5512
activities which developed in the last 3–5 decades, resulting in relatively
high levels of lichen diversity and in moderate or low degrees of air
pollution even in the past years when some areas of the Veneto region
were affected by high levels of SOx pollution (Cislaghi and Nimis, 1997;
Nimis et al., 1991).
Forty percent of the Montello area is managed for extensive
agriculture, including arable fields, hay meadows, and vineyards,
while the remnant 60% is covered by forests. The forest landscape may
be described as a rather continuous black-locust matrix with
interspersed oak-dominated patches. Pure black-locust formations
are, therefore, the main forest type, representing over 90% of the
forested area. They are private forests managed with short-rotation
cycles (15–30 years), mainly for firewood production. Of the original
c. 5000 ha of the oak forest present in the 19th century, only a few
remnant patches (total area 106 ha) remained in the central part of
the hill. They are non-intensively managed with occasional cutting by
the owners. Additional minor formations are chestnut forests and
artificial conifer plantations. Since 2000, Montello has been included
in the Natura 2000 network (IT 3240004).
2.4. Comparison between historical and current lichen data
Information on the epiphytic lichen biota hosted in the past in the
oak forest of Montello was retrieved from two main sources. The
analytical lichen flora of Veneto by Saccardo (1894) includes all the
records which were known for the region until that date. From this
checklist we compiled the records for which it was clearly stated the
provenance from Montello. Further information was retrieved from
the herbarium BASSA which hosts the specimens collected by
Massalongo and Beltramini during their field trip in August 1851
(Nascimbene, 2007). We included only species for which the note on
the collecting site was completed by a precise comment on the habitat
(oak forests). Despite the fact that in a comment to their field work
Massalongo and Beltramini stated that the Montello was a relatively
lichen poor area (Nascimbene, 2007) this dataset is not proposed here
as an exhaustive checklist.
In 2009, we carried out accurate floristic surveys in all (n = 43) the
oak remnant formations (total area 106 ha), checking if the species
sampled between 1851 and 1894 were still present in the remnant
forests. The minimum, mean and maximum patch size was 0.369, 2.47
and 12.6 ha, respectively. We carefully explored all the microhabitats
potentially hosting the species historically recorded, including the
base of the trunks which are often covered by mosses and are
potentially suitable for example for some Peltigera species.
We retrieved the current rarity status of each species included in
the historical list. Nimis and Martellos (2008) have estimated the
rarity of the species at the national level on the basis of: a) number of
samples in the TSB lichen herbarium, b) number of literature records,
and, c) expert judgement. Eight commonness-rarity classes were
used, from extremely rare to extremely common. The ‘extremely rare’
status is given only to taxa known from less than 5 localities in Italy, or
to those that were not mentioned in the literature in the last 50 years.
Recently-described or dubious taxa are excluded from this category.
Extremely rare species are roughly compared to threatened species
(Nimis and Martellos, 2008) according to the IUCN criteria (2001) and
therefore targeted for conservation purposes at the national level.
Nomenclature of lichens follows Nimis and Martellos (2008).
2.5. Comparison between currently available habitats for epiphytic
lichens
A second lichen survey was performed to compare the three main
habitats which are currently available for the establishment of
epiphytic lichen communities: (1) isolated, mainly subneutral-barked
(e.g. Juglans regia L.), trees in agrarian landscapes providing well-lit
conditions, (2) pure black-locust forests providing moderately shaded
conditions, and (3) remnant oak forests not including black-locust
trees, providing shaded conditions and subacidic–acidic substrates.
For each of the three habitat types, six 30 × 30 m plots were selected in
different parts of the hill in order to encompass the variability of each
habitat in the study area (Fig. 1). The minimum distance between
plots of the same habitat was 500 m to reduce spatial dependence.
The borders of the forest plots were at least 50 m far from the forest
edge. Agricultural plots were selected where at least six trees with
DBH N 15 cm could be found within a 900 m2 area. In each plot, 6 trees
with DBH N 15 cm were selected for lichen survey by random
sampling, for a total of 108 trees (36 in each habitat type). In blacklocust and oak forests only black-locust and oak trees were
respectively used for the lichen survey. In agricultural habitats lichens
were mainly surveyed on J. regia L. (70%), and on Prunus avium L.,
Pyrus sp., and Quercus sp. The lichen inventory followed the guidelines
proposed by Stofer et al. (2003) for the Forest BIOTA project, which
are based on the European guidelines for lichen monitoring (Asta
et al., 2002). Lichen diversity was sampled using four standard frames
of 10 × 50 cm as sampling grids, subdivided into five 10 × 10 cm
quadrats, which were attached to the tree trunk at the cardinal points
with the lower side at 100 cm from the ground. All lichen species
inside the frames were listed and their frequency was computed as
the number of 10 × 10 cm quadrats in which the species occurred.
Nomenclature of lichens follows Nimis and Martellos (2008).
2.6. Data analysis
The comparison between historical and current lichen data was
done only qualitatively as the historical records were not precisely
localized. We, therefore, estimated species extinctions at the regional
level.
Concerning the comparison between currently available habitats,
one-way analysis of variance (ANOVA) was applied to test the effect of
habitat type on species richness. The analysis was performed using
the cumulative number of species per plot (sum of the species found
on the six trees) and mean number of species per tree, separately. In
both cases we used the plot as replicate to avoid pseudo-replication.
After the ANOVA, a Tukey's honest significance test for multiple
comparison was applied to detect differences between the three
habitat types (P b 0.05). To test the influence of habitat type on species
composition, ordination methods were applied. The response variable
was the species by plot matrix based on species frequencies. A
preliminary Detrended Correspondence Analysis (DCA) was performed. The largest DCA gradient length, expressed in the standard
deviation (SD) units of species turnover, of the first four DCA axes was
3.23. Thus, the use of linear-based ordination models was appropriate
for these data (Ter Braak and Šmilauer, 2002). First, a Principal
Component Analysis (PCA) was performed to extract the variability
related to species composition. Second, a Redundancy Analysis (RDA)
was performed, using habitat type as a factor. RDA tested if the three
habitat types differed in species composition using a Monte Carlo
permutation test (n = 1000). We ran both PCA and RDA to evaluate
how much variation in species composition data was explained by
habitat type. For these analyses we used the CANOCO package
(Version 4.5, Ter Braak and Šmilauer, 2002).
3. Results
3.1. Historical vs. current lichen records
The historical list included 30 species and 1 lichenicolous fungus
(Sphinctrina turbinata) (Table 1). They were mainly crustose (55%)
and foliose (35%) species reproducing by ascospores (77%), or soredia
(16%). Fifty-five percent of the species had chlorococcoid green algae,
29% cyanobacteria, and 26% trentepholioid algae as photobiont. Five
species were extremely rare at the national level.
J. Nascimbene, L. Marini / Science of the Total Environment 408 (2010) 5506–5512
Table 1
Comparison of species found in the oak forest during the period 1851–1894 and in
2009. For each species the photobiont type and the current rarity status at the national
level are reported according to Nimis and Martellos (2008). Photobiont: Ch = green
algae other than trentepohlia; Cy = cyanobacteria; Tr = trentepohlioid alga.
Species found between 1851 and 1894
2009
Photobiont
Acrocordia gemmata (Ach.) A.Massal.
Arthonia cinereopruinosa Schaer.
Arthonia cinnabarina (DC.) Wallr.
Arthonia elegans (Ach.) Almq.
Arthonia pruinata (Pers.) A.L.Sm.
Arthonia radiata (Pers.) Ach.
Bacidia rosella (Pers.) De Not.
Calicium abietinum Pers.
Degelia plumbea (Lightf.) M.Jørg. & P.James
Flavoparmelia caperata (L.) Hale
Fuscopannaria leucophaea (Vahl) M.Jørg.
Gomphillus calycioides (Duby) Nyl.
Lecanographa amylacea (Pers.) Egea
and Torrente
Lecidea exigua Chaub.
Leptogium corticola (Taylor) Tuck.
Leptogium cyanescens (Rabenh.) Körb.
Lobaria pulmonaria (L.) Hoffm.
Nephroma resupinatum (L.) Ach.
Pannaria rubiginosa (Ach.) Bory
Parmotrema perlatum (Huds.) M.Choisy
Peltigera canina (L.) Willd.
Peltigera horizontalis (Huds.) Baumg.
Peltigera polydactyla (Neck.) Hoffm.
Pertusaria amara (Ach.) Nyl.
Pertusaria hymenea (Ach.) Schaer.
Pertusaria leioplaca DC.
Pertusaria pertusa (Weigel) Tuck.
Pyrenula nitida (Weigel) Ach.
Pyrgidium montellicum (Beltr.) Tibell
Ramalina farinacea (L.) Ach.
Sphinctrina turbinata (Pers. Fr.) De Not.
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Confirmed
Extinct
Extinct
Extinct
Tr
Tr
Tr
Tr
Tr
Tr
Ch
Ch
Cy
Ch
Cy
Ch
Tr
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Confirmed
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Extinct
Ch
Cy
Cy
Ch/Cy
Cy
Cy
Ch
Cy
Cy
Cy
Ch
Ch
Ch
Ch
Tr
Ch
Ch
–
Nationally
rare
+
+
+
+
+
Only two species out of 31 were found again in 2009 (Flavoparmelia
caperata and Parmotrema perlatum). Both species are still common in
Italy. P. perlatum is more frequent in areas with suboceanic climate, such
as the Tyrrhenian side of the peninsula, while in the north it is restricted
to remnant humid forests (Nimis and Martellos, 2008).
3.2. Current lichen diversity
Forty-seven species were found in the 18 plots (Appendix).
Cumulative species richness as well as mean species richness was
highest in agrarian stands and lowest in oak forests, black-locust
forests being in an intermediate position (Fig. 2). Also the overall
number of species increased from oak to black-locust forests and
agrarian stands (16, 19, and 37). Thirteen percent of the species
occurred in all the habitat types, being however more frequent in
black-locust forests and agrarian stands, such in the case of the two
nitrophytic species Candelaria concolor and Phaeophyscia chloantha.
In the PCA ordination (Fig. 3a), the two axes represented 64.3% of the
total variation in species composition (43.4% axis 1 and 20.9% axis 2).
Plots belonging to different habitats were clearly separated, indicating
major differences in species composition. Black-locust forests were in an
intermediate position between oak forests and agrarian stands. In the
RDA ordination, which was constrained to the habitat types (Fig. 3b), the
two axes represent 52.6% of the total variation (38.4% axis 1 and 14.4%
axis 2), i.e. the constrained ordination explained a large proportion of
variation in species composition. A high number of species was related
to agrarian stands. However, each habitat included some associated
species, such in the case of Catillaria nigroclavata and Hyperphyscia
adglutinata for black-locust forests, and Arthonia spadicea, Lepraria
lobificans, and P. perlatum for oak forests. The forest species Dimerella
pineti is shared between black-locust and oak forests.
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4. Discussion
Our study found severe species extinction comparing the available
historical records with those found on the remnant habitats. Amongst
the species historically recorded in native forests, there is a large
incidence of lichens with cyanobacteria and Trentepohlia as photobiont. These lichens, which in Europe have an oceanic–suboceanic
distribution, are considered old-growth specialists extremely sensitive to intensive forest management (e.g. Campbell and Fredeen,
2004; Kuusinen, 1996), and human impact (e.g. Gauslaa et al., 2007;
Geiser and Neitlich, 2007; Richardson and Cameron, 2004; Wolseley
and Aguirre-Hudson, 1997). In Italy, they are mainly restricted to the
western side of the peninsula and to the rainiest areas of the Alps
where they are associated with humid and scarcely disturbed forests
(Nascimbene et al., 2007, 2009; Nimis and Tretiach, 1995).
In our study region, the continuous native oak forest was
transformed into either black-locust formations or arable lands,
while only a few remnant patches are still preserved. According to
Belinchón et al. (2009), destruction and fragmentation of forest
habitats represent one of the most important threats for epiphytic
lichens. The loss of the native habitat, the subsequent invasion by
black locust and the agricultural management, were probably the
most influent factors which affected and modified the composition of
lichen communities (Aragón et al., 2010; Loppi and De Dominicis,
1996; Van Herk et al., 2003), causing the local extinction of most of the
species historically recorded and the establishment of lichen
communities composed by eutrophication-tolerant species. Air
pollution might potentially have contributed to this process but the
relatively high values of lichen diversity found in this area by Nimis
et al. (1991), indicating low impact of SOx pollution, allowed us to
hypothesize that metapopulation processes are likely to be more
important than air pollution in our specific case. This hypothesis is
also corroborated by the fact that some of the species still found in the
remnant oak forests, such as A. spadicea and P. perlatum are sensitive
to air pollution (Nimis, 1993). The historical and current pollution
conditions of our study region are significantly different from that in
central and eastern Europe, where large industrial areas related to
metal and coal mining and transformation developed since the end of
19th century, producing severe impacts on epiphytic lichens and
causing the extinction of many species even in remote forested areas
which were greatly impacted by a long-distance acidic deposition.
Despite high and diffuse anthropization, the landscape of northeastern Italy is still largely dominated by agriculture mixed with small
industries which mainly developed in the last 3–5 decades.
Considering the habitats currently available to epiphytic lichens,
remnant oak forests host lichen communities which clearly differ in
species composition from both agrarian habitats and black-locust
forests. Oak remnants hosted only a few species which were found in
native forests and were the lichen poorest habitat in the study region.
Aragón et al. (2010) have demonstrated that in oak forests, management intensity was the main predictor affecting the total epiphytic
richness, suggesting that our small remnants could be influenced by past
intensive management, resulting in species poor lichen communities.
However, it is noteworthy that these communities were mainly
composed by sub-acidophytic, shading tolerant, and nitrogen intolerant
species (Nimis and Martellos, 2008), indicating that in this habitat the
current level of eutrophication through nutrient deposition from the
surrounding agrarian landscape is still low (Loppi and De Dominicis,
1996; Van Herk et al., 2003). Agrarian habitats were the species richest
habitat. However, they hosted lichen assemblages dominated by lichens
preferring well-lit and dry conditions, most of which are related to
eutrophicated substrates (Nimis and Martellos, 2008). Despite the fact
that eutrophication could cause impoverishment and shifts in species
composition (Aragón et al., 2010; Loppi and De Dominicis, 1996),
several nitrophytic lichens are enhanced by high levels of N deposition,
resulting in relatively species rich communities (Loppi and Nascimbene,
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J. Nascimbene, L. Marini / Science of the Total Environment 408 (2010) 5506–5512
Fig. 2. Results of the one-way ANOVA and Tukey's honest significance test for multiple comparisons. We tested both (a) the cumulative number of species per plot and (b) the mean
number of species per tree. We used the 18 plots as replicates in both analyses. Bars indicate mean with SE while different letters mark significant differences at P b 0.05.
2010). The ecological requirements of the species found in black-locust
forests were similar to those inhabiting agrarian stands. The presence of
a few forest lichens, such as D. pineti, which was mainly associated with
oak remnants, or the rare Agonimia allobata, which was recently found in
the region within riparian forests (Nascimbene et al., 2008), indicates a
potential of these alien formations for hosting specific lichen communities. In this perspective, further research is needed to clarify if there is a
succession during the rotation cycle (see e.g. Rogers and Ryel, 2008),
with a shift from open habitat to forest habitat communities, and if a
prolonged rotation cycle could allow black-locust stands to be suitable
surrogates for richer forest lichen assemblages.
4.1. Management implications
Our study indicated that the loss of the native oak forest with the
subsequent invasion by black locust was probably the major cause of
local extinction of several rare lichen species. The few remnant oak
forests were species poor and mainly composed of common species.
Since oak forests are targeted amongst the habitats of conservation
concern in Europe (e.g. see Council Directive 92/43/EEC), future forest
management should aim at restoring a more natural forest composition,
favouring the spread of oaks and other native trees in lowland systems
which are included in the Natura 2000 network. This might be expected
to produce positive effects on lichen communities, providing more
habitat area and more buffered conditions enhancing the establishment
of forest species. The invasiveness of black locust is the main constraint
to restoring large oak forests in the study area due to its fast vegetative
reproduction. Since an inversion of the ratio between oak and blacklocust forests does not seem realistic even in the long term, it should be
investigated if some changes in the management of black-locust forests
could be introduced to promote more conservation-oriented practices.
These problems are likely to be common also in other lowland and hilly
regions throughout Northern Italy, involving Natura 2000 sites (e.g.
Motta et al., 2009). For this reason, a national strategy for biodiversity
conservation and monitoring of lowlands forests should provide the
framework for local restoration projects.
Fig. 3. (a) PCA ordination diagram of the 18 plots against the first two Principal Components and (b) RDA ordination diagram of lichen species occurring in the 18 plots against the
first two canonical axes constrained by habitat type. In parentheses the explained variation by each axis is reported. In the RDA plot, only species with a fit-range above 10% are
shown. ▲ Categorical variables; ↑ Species. Abbreviations as in Appendix.
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J. Nascimbene, L. Marini / Science of the Total Environment 408 (2010) 5506–5512
Appendix
(continued)
A (continued)
Acknowledgements
Species
The work was economically supported by the Civic Museum of
Montebelluna (Treviso).
Francesco Bortignon (Montebelluna) is thanked for his contribution during the field work, Francesco Ferrarese (Padova) for providing
GIS information, and ARPAV, Centro Meteorologico di Teolo, for
providing climatic information related to the study area (Volpago del
Montello). L.M. was supported by the PRATIQUE project (Enhancements of Pest Risk Analysis Techniques) financed through the EU
Seventh Research Framework Program (FP7) (Contract No. 212459).
We are grateful to the Associate Editor James P. Bennett and to
three anonymous referees for providing useful suggestions and
stimulating comments which improved the clarity and the effectiveness of the manuscript.
Appendix A
Species found in the 18 plots. Frequency in each habitat type is
expressed as the proportion of plots in which the species occurred.
BL = Black-locust forests; O = Oak forests; A = Agrarian. * marks
nitrophytic species, while ° marks shading tolerant species according
to Nimis and Martellos (2008).
Species
Abbreviations Frequency (%)
Agonimia allobata (Stizenb.) P.James°
Anisomeridium biforme (Borrer) R.C.Harris
Arthonia radiata (Pers.) Ach.
Arthonia spadicea Leight.°
Arthrosporum populorum A.Massal.
Caloplaca cerinella (Nyl.) Flagey*
Caloplaca ferruginea (Huds.) Th.Fr.
Caloplaca obscurella (Körb.) Th.Fr.
Caloplaca pyracea (Ach.) Th.Fr.
Candelaria concolor (Dicks.) Stein*
Candelariella efflorescens auct. eur.*
Candelariella reflexa (Nyl.) Lettau*
Candelariella xanthostigma (Ach.) Lettau
Catillaria nigroclavata (Nyl.) Schuler
Cladonia coniocraea (Flörke) Spreng.
Dimerella pineti (Ach.) Vezda°
Evernia prunastri (L.) Ach.
Flavoparmelia caperata (L.) Hale
Flavopunctelia flaventior (Stirt.) Hale
Hyperphyscia adglutinata (Flörke) H.
Mayrhofer & Poelt*
Hypogymnia physodes (L.) Nyl.
Lecania cyrtella (Ach.) Th.Fr.
Lecanora carpinea (L.) Vain.
Lecanora chlarotera Nyl.*
Lecanora sambuci (Pers.) Nyl.
Lecidella elaeochroma (Ach.) M.Choisy
Lecidella pulveracea (Schaer.) P.Syd.
Lepraria incana (L.) Ach.°
Lepraria lobificans Nyl.°
Melanelixia fuliginosa (Duby) O. Blanco. A.
Crespo. Divakar. Essl.. D. Hawksw. &
Lumbsch
Melanelixia subaurifera (Nyl.) O. Blanco. A.
Crespo. Divakar. Essl.. D. Hawksw. &
Lumbsch
Normandina pulchella (Borrer) Nyl.
Parmelia sulcata Taylor
Parmelina tiliacea (Hoffm.) Hale
Parmotrema perlatum (Huds.) M.Choisy
Phaeophyscia chloantha (Ach.) Moberg*
Phaeophyscia hirsuta (Mereschk.) Essl.*
Phaeophyscia orbicularis (Neck.) Moberg*
Phaeophyscia pusilloides (Zahlbr.) Essl.
Physcia adscendens (Fr.) H.Olivier*
agoal
anibif
artrad
artspa
arpopu
calcer
calfer
calobs
calpyr
cancon
caneff
canref
canxan
catnig
clacon
dimpin
evepru
flacap
flafla
hypadg
BL
A
hyphys
leccyr
lecarp
lechla
lesamb
leciel
lecifla
lepinc
leplob
melful
0
16.7
0
0
0
16.7
0
0
0
0
melsub
0
norpul
parsul
partil
parper
phachl
phahir
phaorb
phapus
phyads
O
16.7
0
0
33.3
0
16.7
0
33.3
0
0
0
100
0
16.7
0
0
33.3
0
0
16.7
0
16.7
0
0
0
50
0
100
100
16.7
83.3
50
0
50
50
16.7
50
66.7
16.7
83.3
83.3
0
0
0
33.3
66.7
0
100
0
33.3
0
0
66.7
33.3
0
16.7
0
100
100
0
33.3
0
33.3
0
33.3
0
50
16.7
33.3
0
83.3
0
0
33.3
0
16.7
0
100
33.3
0
16.7
66.7
33.3
0
33.3
0
16.7
0
0
66.7 100
16.7
66.7
16.7 100
0
16.7
83.3 100
0
83.3
0
0
50
16.7
0
0
0
0
(continued on next page)
Abbreviations Frequency (%)
BL
Physcia clementei (Turner) Maas Geest.
Physconia distorta (With.) J.R.Laundon*
Physconia perisidiosa (Erichsen) Moberg
Pseudevernia furfuracea (L.) Zopf v. furfuracea
Punctelia borreri (Sm.) Krog
Punctelia subrudecta (Nyl.) Krog
Xanthoria parietina (L.) Th.Fr.*
phycle
phydis
phyper
psefur
punbor
punsub
xanpar
A
0
50
0
16.7
0
16.7
0
33.3
16.7
50
0
33.3
16.7 100
O
0
0
0
0
16.7
0
0
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