Science of the Total Environment 505 (2015) 315–328
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Science of the Total Environment
journal homepage: www.elsevier.com/locate/scitotenv
Black locust—Successful invader of a wide range of soil conditions
Michaela Vítková a,⁎, Jaroslav Tonika b, Jana Müllerová a
a
b
Institute of Botany, Academy of Sciences of the Czech Republic, 25243 Průhonice, Czech Republic
Faculty of Science, Charles University, Institute for Environmental Studies, Benátská 2, 12801 Praha 2, Czech Republic
H I G H L I G H T S
G R A P H I C A L
A B S T R A C T
• We provided an overall assessment of
black locust soil conditions.
• Black locust tolerates extremely diverse
soil physical–chemical properties.
• Black locust seems to be limited by
water supply and soil aeration.
• The most common are young soils
(Cambisols, Leptosols and Arenosols).
• Species composition in BL stands was
mostly affected by soil reaction.
a r t i c l e
i n f o
Article history:
Received 6 June 2014
Received in revised form 29 September 2014
Accepted 29 September 2014
Available online xxxx
Editor: Charlotte Poschenrieder
Keywords:
Bedrock
Nitrification
Physical–chemical soil properties
Plant invasion
Robinia pseudoacacia
Soil type
a b s t r a c t
Black locust (Robinia pseudoacacia, BL), a species native to North America, has successfully invaded many types of
habitats over the world. This study provides an overall assessment of BL soil conditions to determine the range of
physical–chemical soil properties it can tolerate. 511 BL stands (for the soil types) and 33 permanent plots (for
the soil chemistry) were studied in the Czech Republic. Relationships among different environmental variables
(physical–chemical soil properties, vegetation characteristics and habitat conditions) were investigated and variables with the highest effect on species composition were detected. The results were compared with data in the
literature for other parts of the secondary and native distributions of this species. This assessment showed that BL
is able to tolerate extremely diverse soil physical–chemical conditions, from extremely acid to strongly alkaline,
and from medium to highly base saturated soils with a gradient of different subsurface stoniness. Soil nitrate, N
mineralization and nitrification rates also varied considerably and the concentrations of exchangeable phosphorus and ammonium were consistently low. N mineralization rate, incubated inorganic nitrogen and nitrates were
positively correlated with base saturation and cation exchange capacity. The most common soil types were young
soils (Cambisols, Leptosols, Arenosols, and coarsely textured Fluvisols). BL seems to be limited by water supply
and soil aeration and prefers well aerated and drained soils, and tolerates desiccation but avoids compact soils
and areas where the soils are frequently waterlogged. On steep slopes, BL was less vigorous, stunted and less
competitive. By contrast, the tallest BL trees were found on sandy soils in a flat landscape. Number and share
of nitrophytes in the herb layer were positively related to basic bedrock, soil reaction and N–NO3/N ratio. Soil
reaction was determined as the most important environmental characteristic explaining the variability in BL
species composition in the Czech Republic.
© 2014 Elsevier B.V. All rights reserved.
Abbreviations: Ass, association; BL, black locust; BS, base saturation; C, total carbon; CCA, canonical correspondence analysis; CEC, cation exchange capacity; Corg, organic carbon; PCA,
principal component analysis; Pex, exchangeable phosphorus; r.s, Spearman's nonparametric correlation coefficient.
⁎ Corresponding author. Tel.: +420 271015241; fax: +420 271015105.
E-mail addresses: michaela.vitkova@ibot.cas.cz (M. Vítková), jaroslav.tonika@gmail.com (J. Tonika), jana.mullerova@ibot.cas.cz (J. Müllerová).
http://dx.doi.org/10.1016/j.scitotenv.2014.09.104
0048-9697/© 2014 Elsevier B.V. All rights reserved.
316
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
1. Introduction
Black locust (Robinia pseudoacacia), a nitrogen fixing tree belonging to the family Fabaceae, was introduced from its native range in
North America (Fowells, 1965; Huntley, 1990) to other continents
and currently is naturalized in Europe, temperate Asia, Australia,
New Zealand, northern and southern Africa and temperate South
America (Weber, 2003). In its native range, BL is listed as a component of mixed mesophytic forests and also readily colonizes open
sites created by fire, floods, logging or storms (Boring and Swank,
1984a). The degree of invasion of different habitats in Europe differs
(Chytrý et al., 2008). Chytrý et al. (2005) state that R. pseudoacacia is
one of the top 10 neophytes with the broadest habitat range. It
spreads mostly vegetatively by means of its aggressive root and
trunk coppice shoots, whereas the seedlings are successful only on
bare soil. The most invaded natural habitats include thermophilous
grasslands (Kleinbauer et al., 2010; Vítková and Kolbek, 2010),
sandy soils, shrubbery and azonal forests, such as thermophilous
oak, dry acidophilous oak, dry pine (Vítková and Kolbek, 2010),
maple-lime (Kleinbauer et al., 2010), chestnut and riparian forests
(Brus, 2006; Motta et al., 2009; Benesperi et al., 2012; GonzálezMuñoz et al., 2013), urban–industrial wastelands, fallow lands, disturbed traffic corridors and at burnt sites (e.g. Dzwonko and Loster,
1997; Kim and Lee, 2005; Řehounková and Prach, 2008; Yüksek,
2012; Kowarik et al., 2013). Both climate change (Kleinbauer et al.,
2010) and planting for forestry or landscaping (Kowarik, 2003) are
likely to increase its distribution and possibly also enlarge the
range of habitats it is able to colonize, including Central European
zonal forests (Essl et al., 2011).
BL causes homogenization of the tree layer and creates specific
stands, highly different from autochthonous plant communities (e.g.
Wendelberger, 1954; Montagnini et al., 1991; Peloquin and Hiebert,
1999; Von Holle et al., 2006; Taniguchi et al., 2007; Kolbek and
Jarolímek, 2008; Vítková and Kolbek, 2010; Benesperi et al., 2012;
Sitzia et al., 2012). The strong effect of BL on native vegetation is probably caused by increased nutrient availability associated with the nitrogen fixing ability of the symbiotic Rhizobium bacteria (37 strains)
occurring in BL root nodules (Batzli et al., 1992; Ferrari and Wall,
2007). Symbiotic fixation is an important input for the nitrogen cycle
in BL stands, more important than litter mineralization or other
sources (e.g. Liu and Deng, 1991; Tian et al., 2003; Williard et al.,
2005). In its native range (southern Appalachian forests) it can fix
33 to 75 kg N ha− 1 year− 1, with a particularly high capacity for N2
fixation in early to intermediate stages of secondary succession
(Boring and Swank, 1984b). Symbiotic fixation recorded in its secondary range is even greater, 110 kg N ha−1 year−1 by four-year-old trees
in Austria (Danso et al., 1995) and 112.3 kg N ha−1 year−1 by 25-yearold south-facing stands in Central Korea (Noh et al., 2010). According
to Liu and Deng (1991), the main factors determining nitrogen fixation
are soil acidity and available phosphorus.
Occurrence of nitrogen fixing trees in forest ecosystems results in
subsequent increase in the soil nitrogen pool, nitrification and net Nmineralization rates and higher availability of mineral forms of nitrogen
−
(ammonium, NH+
4 and nitrate, NO3 ) in both soil and solution (e.g.
Binkley et al., 1982; Van Miegroet and Cole, 1984; Montagnini et al.,
1991; Montagnini and Sancho, 1994). Enriched level of soil nitrogen is
not only a result of release from decaying N-rich BL leaves and roots,
but also from root exudates, which contain 1–2% of recently fixed N
(Uselman et al., 1999; Tateno et al., 2007). High rates of soil nitrification
can result in a decrease in pH values of litter and topsoil and potentially
a greater leaching of Ca, Mg, K, Na and PO4–P ions from the soil (Van
Miegroet and Cole, 1984). However, some authors, e.g. Montagnini
and Sancho (1994) or Rice et al. (2004), have not confirmed the
supposed acidification effect of nitrification. The allelopathic potential
of BL has only been recorded in the laboratory (Nasir et al., 2005;
Csiszár, 2009).
The first recorded BL plantation in the Czech Republic was in 1785
(Nožička, 1957). It was planted mainly in former pastures on steep
eroded hillsides along rivers (especially Vltava—Fig. 1, Berounka, Sázava
and Dyje) in order to stabilize sandy soils, aeolian sands and coarse
fluvial deposits (Elbe lowland, Fig. 2, and parts of South Moravia) and
ameliorate poor soils, and around transport corridors (Kolbek et al.,
2004). Currently, it covers approximately 12,000 ha, i.e. 0.46% of the
total forested area in the Czech Republic and occurs in most areas of
the country below 500 m a.s.l. The largest stands are concentrated in
the warmest part of the country, preferring south-facing slopes of
30–40° (Vítková et al., 2004). As a source of fast growing and valuable
wood ameliorating poor soils it is widely used in forest plantings,
resulting in rapid spread to natural systems (covering 0.2% of Czech
Republic “NATURA 2000” sites). From phytosociological point of view,
there are four types of Robinia stands with different soil conditions:
(1) species-rich nitrophilous stands growing on alkaline to acid bedrocks (ass. Chelidonio majoris-Robinietum pseudoacaciae) (Sádlo et al.,
2014; Fig. 3); (2) species-poor grassy stands—tall BL forests with
straight trunks (ass. Arrhenathero elatioris-Robinietum pseudoacaciae)
on strongly acid quaternary deposits (Sádlo et al., 2014; Fig. 2);
(3) open and mesic stands with the herb layer dominated particularly
by Poa nemoralis (ass. Poo nemoralis-Robinietum pseudoacaciae) on
upper and middle slopes on siliceous bedrock in deep river valleys
(Sádlo et al., 2014); and (4) dwarf and shrubby stands on thermophilous rocky slopes (ass. Melico transsilvanicae-R. pseudoacaciae;
Sádlo et al., 2014; Fig. 1).
The adaptability of BL to different habitat conditions and an absence
of serious natural enemies in its secondary range makes it an economically attractive tree species, especially for short-rotation energy plantations (e.g. Grünewald et al., 2009; Rédei et al., 2010) and soil
reclamation (e.g. Kim and Lee, 2005; Qiu et al., 2010; Yüksek, 2012).
However, regular silvicultural treatment is needed to maintain the
short-rotation plantation productivity because of relatively low production of litter and periodic removal of organic matter (Vasilopoulos
et al., 2007). Some of the economic benefits of BL, such as its vitality,
excellent sprouting ability, abundant production of seed and improvement of soil conditions by nitrogen fixation, become a problem after
the plantation is abandoned. Stumps of harvested trees resprout rapidly
(e.g. Krízsik and Körmöczi, 2000) and eradication with the aim of restoring original plant communities is very difficult, costly and timeconsuming (e.g. Hruška, 1991; Peloquin and Hiebert, 1999; Halassy
and Török, 2004; Böcker and Dirk, 2004, 2007; Malcolm et al., 2008;
Yong-Chan et al., 2009; Vítková, 2011; Ivajnšič et al., 2012; Skowronek
et al., 2014). Natural succession of abandoned BL plantations towards
the natural communities is slow; Vasilopoulos et al. (2007) did not record any succession towards the nearby natural riparian forests even
after 14 years. This is in marked contrast to the BL native range, where
after 15–30 years BL is replaced by more competitive tree species
(Boring and Swank, 1984a).
Although BL is the second most widely planted woody species in
the world (Keresztesi, 1988), comprehensive information on its soil
conditions is missing. From its native range, the data are rare and
do not cover the range of environmental variability, coming solely
from the Coweeta LTER site (southern part of the native range;
Boring and Swank, 1984a,b; Montagnini et al., 1986; White et al.,
1988; Montagnini et al., 1989; Montagnini et al., 1991). Although
there are more data available from BL secondary range (e.g. Dzwonko
and Loster, 1997; Šimonovič et al., 2001; Noh et al., 2010; Yanna et al.,
2013), they usually come from isolated unevenly distributed sites
only. Our specific objectives were to (1) assess the overall environmental variability of soil types invaded by R. pseudoacacia in the traditional Central European landscape (Czech Republic); (2) determine
the range of physical–chemical soil characteristics tolerated by BL in
the Czech Republic; (3) evaluate the most important characteristics
influencing species composition and amount of nitrophytes in BL stands
and (4) compare our findings with the literature data from the native
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
317
Fig. 1. Rocky habitat invaded by Robinia pseudoacacia in the valley of the Vltava River (Vítková, May 2012).
and other parts of the secondary range to detect similarities in BL soils
in different parts of the distribution range.
2. Methods
2.1. Study area
Due to its history and geomorphology, the Czech Republic can be
considered as an example of traditional Central European landscape.
In our study, 33 permanent plots each with an area of 250 m2 were
established in 1997–1999 (Fig. 4) in particular types of vegetation
(based on a syntaxonomical approach; Vítková and Kolbek, 2010), covering the diversity of bedrocks, soil types and different habitats
(Table 1). On each plot, phytosociological relevé using a seven-grade
scale of abundance and dominance (Braun-Blanquet (1964) was recorded. Nitrophilous species were determined using Ellenberg nitrogen
indicator value six or higher (Ellenberg et al., 1991).
The annual temperature ranged between 6.6 and 9.1 °C, annual
precipitation 494–604 mm (moderate Atlantic-continental climate),
altitude 150–380 m a.s.l. and slope 0–50° (Table 1). The plots were selected in unmanaged successionally mature (at least 40 years old) BL
stands growing in open landscape or suburban areas. Most of the sites
originated as spontaneous BL invasion into natural communities, with
about 20% of the stands self-regeneration from plantations established
at the end of the 19th to the middle of the 20th century. None of the
forests studied were subjected to recent forest management.
2.2. Soil characteristics
At each of the 33 permanent plots a test pit was dug, soil profile was
described and samples of bedrocks, soils and lumps of soil from A and B
horizons were collected. Lumps of soil were impregnated with retin
Epofix-Struers both before and after sectioning. The surface of each
thin section was ground down using carborundum (600). Thin sections
of soil were prepared by the Czech Geological Survey and used to
Fig. 2. Grassy black locust stand on Arenosol and aeolian sand bedrock (Vítková, June 2011).
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M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
Fig. 3. Nitrophilous black locust stand on Cambisol and granodiorite bedrock at the base of slope (Vítková, June 2013).
supplement knowledge on the petrological composition of rocks and
physical–chemical properties of the soil.
Soil was sampled twice, at the end of the growing season (after a hot
and dry summer when legumes mature, which occurred in the second
half of September 1999) and at the beginning of the growing season
(after a cold and wet spring, before black locust bud burst, which occurred in the second half of April 2000). Each 1 kg sample was collected
separately for A and B horizons from five randomly located points. Fresh
soil was immediately sifted in the field through a 2 mm sieve; fine earth
was transported to the laboratory in a cooler and used to determine am−
monium (NH+
4 ) and nitrates (NO3 ). Other soil analyses (pH, base saturation, exchangeable phosphorus, carbonate concentration, total carbon
and nitrogen, and net N mineralization and nitrification rates) were
carried out on air-dried fine earth, some only for one growing season
(see Table 2). Unsifted fresh soil samples in the field were taken with
the shovel into the plastic bags to assess the subsurface stoniness and
into the closable aluminous desiccant dishes to determine gravimetric
soil moisture and dry matter according to ISO 11465 (1993). Stoniness
was calculated in percentage as the ratio of the weight of desiccated
skeleton (inorganic particles larger than 2 mm) and total weight of
dry soil sample. Gravimetric soil moisture was expressed also in percentage as the mass of water per unit mass of dry soil (105 °C). Dry matter is dry soil residue after drying at 105 °C, expressed as a percentage of
soil dried in the air.
Active pH was measured according to McLean (1982). Methods of
Moore and Chapman (1986) were used to determine base saturation
Fig. 4. Black locust stands studied in the Czech Republic.
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M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
Table 1
Habitat characteristics of black locust stands studied in the Czech Republic.
Number
Type
Syntaxon
Soil type
Bedrock
Altitude
Slope
Aspect
Annual
precipitation
Annual
temperature
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-rich nitrophilous
Species-poor grassy
Species-poor grassy
Species-poor grassy
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Open and mesic
Dwarf and shrubby
Dwarf and shrubby
Dwarf and shrubby
Dwarf and shrubby
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Chelidonio-Robinietum
Arrhenathero-Robinietum
Arrhenathero-Robinietum
Arrhenathero-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Poo-Robinietum
Melico-Robinietum
Melico-Robinietum
Melico-Robinietum
Melico-Robinietum
Eutric leptosol
Eutric leptosol
Rendzic leptosol
Rendzic leptosol
Rendzic leptosol
Calcic leptosol
Cambic leptosol
Arenosol
Lithic cambisol
Dystric cambisol
Dystric cambisol
Fluvisol
Fluvisol
Arenosol
Arenosol
Arenosol
Typic leptosol
Typic leptosol
Cambic leptosol
Cambic leptosol
Cambic leptosol
Lithic cambisol
Lithic cambisol
Lithic cambisol
Lithic cambisol
Lithic cambisol
Dystric cambisol
Arenic cambisol
Haplic luvisol
Lithic leptosol
Lithic leptosol
Lithic cambisol
Chernozem
Spilite
Basalt
Diabase
Limestone
Olivine basalt
Calcareous sandstone
Paleozoic schist
Quarzite sandstone
Calcareous sandstone
Quarcite
Quartz diorite
Carbon-Permian sandstone
Calcareous sandstone
Eolian sands
Eolian sands
Eolian sands
Proterozoic schist
Amphibole schist
proterozoic schist
Proterozoic schist
Amphibolite
proterozoic Schist
Phylite
Syenodiorite
Amphibole schist
Amphibole schist
Lydite
Proterozoic schist
Basalt
granodiorite
Spilite
Quartz diorite
Olivine nephelinite
310
360
285
315
270
240
220
210
240
240
280
330
230
150
150
190
290
200
210
280
300
240
350
310
300
350
250
290
380
330
260
300
260
30
35
30
30
10
40
35
30
30
20
25
30
20
0
0
0
45
35
50
35
40
45
35
30
40
35
30
35
20
45
30
30
25
203
158
158
180
135
180
158
113
203
158
158
225
180
–
–
–
113
158
270
270
135
338
225
203
158
113
113
203
203
90
203
158
203
541
539
495
511
532
530
496
513
535
494
566
577
508
498
498
507
573
524
498
511
625
520
586
574
555
604
482
527
580
574
497
566
557
7.9
7.5
9.0
8.4
7.8
8.3
9.1
8.2
8.2
9.0
8.5
7.6
8.4
8.6
8.6
8.3
8.0
8.9
8.4
8.8
7.7
8.8
7.8
8.5
8.4
8.1
8.9
8.3
6.6
8.5
8.5
8.5
7.5
by exchangeable cations (Ca2+, Mg2+, H+, Al3 +) using titration and
atomic absorption spectroscopy (AAS). Exchangeable phosphorus
(Pex) was measured photometrically using a UV–vis Spectrometer
Unicam 400 (Olsen, 1982). Total carbon (C) and nitrogen
(N) content were analyzed according to Monar (1972) with combustion of soil followed by determination of oxides. Carbonate concentration was determined by following ISO/DIS 10693 (1995).
Organic carbon (C org ) was calculated as the difference between
total carbon and carbonate carbon. Nitrogen mineralization and nitrification were measured by incubating in aerobic laboratory incubators for 28 days following ISO 14238 (1997), and after
moistening (3 ml of water added to each 10 g of sample) and preincubation for one week at 28 °C. The results of soil analyses were
converted to dry matter to eliminate the different water content
among soil samples. Cation exchange capacity (CEC), base saturation (BS), and C org /N, N/P ex , N–NO 3 /N and N–NH 4 /N ratios were
calculated from the results of the soil analyses according to
Sparks et al. (1996). Nitrification tests were carried out in the Research Institute for Soil and Water Conservation (VÚMOP) and
other laboratory analyses in the Analytic Laboratory of the Institute
of Botany CAS.
2.3. Soil types
Assessment of the main soil types, where BL is naturalized, was
based on the analysis of 511 different BL stands in the Czech Republic.
310 sites were field surveyed by the authors in 1998 to 2013, describing
habitat characteristics including soil type, vegetation and recent
management (unpubl.). The resting 201 stands were excerpted from literature (Svobodová, 1952; Blažková, 1961; Větvička, 1961; Sofron,
1964; Němec, 1981) and Czech National Phytosociological Database
(Chytrý and Rafajová, 2003), and their soil types derived from soil
maps at a scale of 1:50 000 (Czech Geological Survey). The nomenclature of soil types follows IUSS (2006).
2.4. Data analysis
Data was evaluated using statistical packages NCSS 6.0 and Canoco 5
(Šmilauer and Lepš, 2014). Inter-correlations between two measured
soil characteristics were determined by calculating Spearman's nonparametric correlation coefficient (r.s.). Changes of soil characteristics
with the soil depth were tested using paired two-sample t-tests. An
unconstrained ordination method was used to detect relationships
among habitat conditions (altitude, slope, aspect, annual precipitation,
annual temperature, soil type, bedrock), vegetation characteristics
(syntaxonomical unit, tree height, cover of tree, shrub and herb layer,
total number of species, number of nitrophytes, and share of
nitrophytes cover to total cover of the herb layer) and physical–chemical soil properties (see Chapter 2.2) in multidimensional space. These
55 variables studied in 33 permanent plots were used as response
data. Most of them were quantitative but soil type, bedrock, aspect
and syntaxonomical unit were used as factors. Aspects between 135
and 225° (from SE to SW) were coded as “warm”, the others as “cold”.
Ion concentrations with differences larger than one order of magnitude
were log transformed before Canoco analysis. Because response data
were not compositional, principal component analysis (PCA) with scaling and standardizing was performed. To decide which of the soil properties describe best the variability in species composition of BL stands,
we used constrained ordination with phytosociological relevés as
response data and soil properties as explanatory variables. Due to long
gradient (4.5 SD units) canonical correspondence analysis (CCA) with
forward selection was chosen. The Monte Carlo permutation test with
999 unrestricted permutations was used to test the significance of
the constrained ordination model. Comparison of results between
constrained and unconstrained ordination applied to the same response
320
Table 2
Physical–chemical characteristics of the soils in black locust stands growing in different parts of the world.
Native range
Secondary range
USA1,2
Poland3
Number of plots
Annual average
temperature [°C]
Annual
precipitation [mm]
Altitude [m a.s.l.]
Stoniness [%]
Soil moisture [%]
pH (KCl)
pH (H2O)
Ca2+ [mg/kg]
2+
Mg
Al
3+
[mg/kg]
[mg/kg]
H+ [mg/kg]
CEC [meq/100 g]
BS [%]
NO3–N [mg/kg]
NO3–N/N [%]
NH4–N [mg/kg]
Croatia5
Korea6
China7,8
Czech Republic
Min
7
8
September
May
–
–
October /July
3
13
1
7.7
3
9.5
1
10.1
2
12.2
2
15.4/8.8
1810
686
600–650
950
1344
1000
242
April
Min
Max
Max
September April
Average (s.d.)
September April
33
7.5
9.1
8.2
1106.5/505
482
625
542
447/1431
150
380
270
110–135
100–120
–
–
–
25–26
–
–
–
27.8–29.3 25.9/–
25
–
3.0
26
–
29.3
0
0
7.0
–
–
2.1
90
–
80
–
133.1 86.1
0–10 –
11–25 –
0–10 29.72
–
–
3.0
140–230
–
–
–
11–25 28.52
–
–
–
–
–
–
–
4.8
1.2
118.4 20.3
0–10
11–25
0–10
11–25
0–10
–
–
5.6
–
230
3.6–5.2
4.0–5.2
4.0–5.2
4.6–4.9
–
4.2
4.9
5.4
5.7
–
–
–
4.3–4.6
–
–
–
–
4.6/8.8
–
–
3.6
4.0
4.0
4.6
230
5.2
5.2
8.8
5.7
230
2.8
3.1
3.4
3.65
–
3.0
3.5
3.5
4.2
264
7.6
8.0
7.9
8.3
–
7.2
7.6
7.6
8.1
20,956
31 (23)
28 (15)
30.8
(27.2)
24.0
(29.2)
4.7 (1.4)
4.7 (1.6)
5.2 (1.3)
5.5 (1.4)
–
11–25 307/322
–
–
–
–
–
–
–
–
112
–
15,828
–
0–10
11–25
0–10
11–25
0–10
11–25
0–10
11–25
0–10
11–25
0–10
11–25
0–10
8.8
–
–
–
–
–
1.2
–
–
–
35.5
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
11.0–15.1
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
8.8
–
–
–
–
–
1.2
8.8
–
–
–
–
–
15.1
–
–
35.5
–
–
–
–
35.5
–
–
–
–
–
–
–
–
–
–
–
–
2.1
0.1
0.02
13
29
0
0
0
0
5.7
4.0
31
18
0.0
0.0
0.00
–
–
–
–
–
–
–
–
–
–
30.1
13.4
0.71
1945
2438
2363
1465
44.9
15.1
113.3
99.2
100
100
203.4
12.1
0.81
–
–
–
–
–
–
–
0.00
0.00
0.93
0.65
13.6
–
–
–
–
–
–
–
–
–
13.6
–
13.6
–
1.7
0.9
1.5
1.3
45.2
9.0
33.7
28.6
–
–
–
–
–
–
–
–
–
–
12.1 (9.2)
5.6 (4.3)
0.18
(0.16)
0.27
(0.27)
7.8 (8.0)
3.5 (1.9)
–
–
–/5.8/5.62
5.72
735/773
149/151
86/82
–
–
55.0/62.9
52.4/62.6
10.7/11.7
7.67/8.73
47/45
21/27
4.5/3.3/11.72
1.6/0.9/3.32
0.37/0.23/
0.352
11–25 0.27/0.14/
0.232
0–10 3.9/2.6/4.92
11–25 3.31/2.18/
1.682
#
September
–
–
15.0 (17.3)
7.3 (5.1)
4.5 (1.3)
4.8 (1.6)
5.8 (1.3)
5.6 (1.5)
4840
(4888)
3600
(4396)
404 (413)
474 (664)
523 (648)
386 (455)
7.5 (8.8)
5.6 (4.6)
34.1 (25.0)
26.8 (25.0)
74 (24)
69 (29)
24.3 (34.4)
3.4 (3.2)
0.34 (0.20)
0.16 (0.16)
6.8 (6.2)
4.6 (6.3)
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
Depth North Carolina
(cm) March1/July1/
Average2
Secondary range
Slovakia4
NH4–N/N [%]
N [%]
C–CO2−
[%]
3
C [%]
Corg [%]
Pex [mg/kg]
N/Pex
Nitrification
[mg/kg/28 days]
Ammonification
[mg/kg/28 days]
N mineralization
[mg/kg/28 days]
#
1
2
3
4
5
6
7
8
0.32/0.18/
0.152
11–25 0.57/0.35/
0.112
0–10 0.12/0.14/
0.332
11–25 0.06/0.06/
0.152
0–10 –
–
–
–
–
–
–
–
0.03
0.02
0.28
0.23
–
–
–
–
–
–
–
0.02
0.03
0.77
1.61
0.10
0.06–0.30
0.25
0.08–0.24 0.47/0.70
0.06
0.70
0.18
0.18
2.61
2.55
–
0.04–0.13
–
–
–
0.04
0.13
0.06
0.09
2.14
0.39
–
–
–
–
–
–
–
0.00
0.00
0.44
1.44
11–25 –
–
–
–
–
–
–
–
0.00
0.00
1.02
2.92
1.05
–
3.50
–
5.75/–
1.05
5.75
2.16
1.96
27.64 31.31
11–25 2.232
–
–
0.40
–
–
0.40
0.40
0.68
1.18
19.55 5.07
0–10
2.38/2.63
–
0.58–5.00
–
1.96–4.12 –/5.90
0.58
5.9
1.94
1.96
27.64 31.31
11–25 1.15/1.26
–
0.05–0.16
–
–
0.05
0.16
0.55
1.18
19.07 5.07
0–10
10.5
1.4–16.7
13.2
17.2–24.5 12.2/8.4
1.4
24.5
9.5
10.3
15.3
15.9
–
35.6
–
–
–
–
1.0–1.3
–
–
–
–
–
–
–
–
–
–
–
–
9.1–18.4
–
–
–
–
–
–/2.5
–
–
–
–
1.0
2.5
–
–
–
–
1.3
35.6
–
–
–
–
4.0
–
–
–
–
–14.2
8.2
1.5
0.2
36
27
–
15.0
–
–
–
–
299.6
15.6
110.9
45.2
3259
9556
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–71.6
–
–
–
–
108.9 –
–
–
–
–
–
–
–
–13.8
–
327.1 –
–
–
–
–
–
–
–
–
–
–
0–10
11–25
0–10
11–25
0–10
11–25
0–10
16–30
0–10
0–10
4.33
2
19.7/18.3/
13.62
20.3/21.2/15.22
10.7/9.8
5.7/3.9
114/145
102/159
34.5/34.3/
36.02
12.0/10.2
–3.62
34.9/30.9/
33.02
16–30 12.2/7.4
Blocks of rock or bedrock are not included in stoniness.
Montagnini et al. (1986)—2 localities (Coweeta hydrologic laboratory), March/July 1982.
Montagnini et al. (1989)—1 locality (Coweeta), average values of soil properties from April to October 1984.
Dzwonko and Loster (1997)—1 locality (Kraków region).
Šimonovič et al. (2001)—3 localities (Záhorie).
Vrbek and Pilaš (2011)—1 locality (Đurđevački pijesci).
Noh et al. (2010)—2 localities (Seoul).
Yanna et al. (2013)—1 locality (Zijin Mountain).
Wang et al. (2012)—1 locality (Zhifangghou, Loess Plateau).
–
–
0.10
(0.07)
0.17
(0.14)
0.89
(0.60)
0.33
(0.40)
0.06
(0.10)
0.14
(0.27)
10.97
(7.40)
3.72
(3.65)
10.91
(7.43)
3.58
(3.61)
12.3 (1.6)
11.5 (2.4)
–
–
–
–
81.2
(66.0)
–
–24.4
(34.6)
56.8
(67.1)
–
0.10 (0.06)
0.26 (0.36)
0.71 (0.52)
0.21 (0.08)
0.07 (0.25)
0.21 (0.70)
8.84 (7.12)
2.57 (1.27)
8.78 (7.12)
2.35 (1.15)
12.0 (1.4)
11.3 (1.7)
20.9 (20.9)
11.4 (12.1)
691 (689)
941 (2248)
–
–
–
–
–
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
Corg/N or C/N
0–10
321
322
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
3%
2%
3% 2%
7%
Cambisols
3. Results and discussion
Leptosols
3.1. Soil types in Central European BL stands
Haplic Luvisols
7%
41%
Fluvisols
Chernozems
Regosols and
Arenosols
Albeluvisols
35%
others
Fig. 5. The most common soil types under the black locust stands growing in the Czech
Republic.
data (phytosociological relevés) allowed to evaluate percentage efficiency of constrained axis.
2.5. Literary comparison
All available data on BL physical–chemical soil characteristics published in international scientific journals or national journals in English
or at least with an English abstract or summary were collected using
Web of Science and Google Scholar. These were used for comparison
of soil conditions in different parts of BL native and secondary range.
In Central Europe, BL grows mostly on young soils in which horizon
differentiation is beginning (86%; Fig. 5). Most frequent are Cambisols
(41%; Fig. 5) with a weak, mainly brownish Cambic subsurface B horizon, which are the most common soils in the Czech Republic (45%).
The second most abundant soils are azonal Leptosols (35%; Fig. 5) on
river valley slopes in which the A horizon is either very shallow on
hard (Lithic Leptosols) or highly calcareous material (Rendzic and
Calcaric Leptosols), or deeper but extremely gravelly and/or stony
soils. Their shallowness or stoniness implies low water holding capacity
and serious limitation for vegetation cover. These conditions cause
shrubby growth form of BL (up to 5 m tall) and decline its competitiveness in favour of drought-adapted native shrubs such as Crataegus sp.,
Prunus spinosa and Rosa sp. Erosion control was therefore the main
reason for BL plantations (Fig. 1). At the bases of slopes, BL stands are
usually found on azonal Fluvisols with a weak brown AC-profile (7%;
Fig. 5). The last category of young soils under BL stands is Regosols
and Arenosols on sandy soils (Fig. 2; 3% in Fig. 5). The later soil type
proved to be the most suitable for BL growth (Fig. 6). Arenosols in the
flat landscape of the Elbe River lowlands are often invaded by speciespoor grassy BL forests with the BL tree height up to 30 m (Fig. 2).
All of the above mentioned soil type groups are well drained. The
long-term filling of soil pores with stagnant groundwater is not usual.
The deeper soils under BL stands are Haplic Luvisols (7%), Chernozems
(3%) and Albeluvisols (3%; Fig. 5).
3.2. Physical–chemical properties in Central European BL stands
1.0
3.2.1. Stoniness
The range of subsurface stoniness varies from 0 to 90%, with less
than 10% on eolian sand, sandstone and basaltoid bedrock. The most
frequent are gravel and stony soils with more than 50% on Ordovician
and Proterozoic schists, amphibolitic schists and spilites. Rocky outcrops
are common.
NO3 Slope
N_inorg_incub.
Stoniness N C Corg
NH4 Open and mesic stands
NH4incub. Al3+
H+
NO3incub.Mineralization
CEC BS
Ca2+
Mg2+
Leptosol
N-NO3/N
Aspect_warm
pH(KCl)
pH(H2O)
C-CO3
Nitrophytes_ Nr
Basic_bedrock Species-rich nitrophilous stands
Nitrophytes_Ratio
Acid_bedrock
Tree_height
Arenosol
Plain
Species-poor grassy stands
A_Horizon
-1.0
E1
-1.0
3.2.2. Gravimetric soil moisture
In the Czech Republic, BL stands occur mostly on dry or drying out
soils, with the two exceptions—a) Eutric Leptosols on spilite with
biogenic aggregate formation caused by high edaphon activity and
b) Dystric Cambisols at the base of lydite outcrops with a thick Oh-horizon, where the gravimetrically determined soil moisture content of fine
earth was as high as 86.1% and 70.8% respectively even at the end of a
hot and dry summer. After excluding these exceptions, the average
soil moisture in A-horizon was 11%. The differences between localities
were more marked after a wet spring when average soil moisture was
up to 31%. The driest soils were recorded on weathered parent rocks
with sandy eluvium (granodiorites, quartz diorites, syenodiorites) and
eolian sands. After wet and cold springs, sands with dust particles
between quartz grains were able to retain water. BL does not grow on
wet soils in which the water has stagnated for a long period; however
on such Arenosols in the Elbe River lowland, BL stands survived even a
centennial flood. The only effect of this flood in 2003 was an increase
in the cover of the nitrophilous species Chelidonium majus (probably
due to nutrient enrichment).
1.0
Fig. 6. PCA ordination diagram for 55 environmental variables—habitat conditions (altitude, slope, aspect, annual precipitation, annual temperature, soil type, bedrock), vegetation characteristics (syntaxonomical unit, tree height, cover of tree, shrub and herb layer,
total number of species, number of nitrophytes, and percentage of nitrophytes in herb
layer) and physical–chemical soil properties detected on 33 permanent plots. 35 bestfitting characteristics are shown. First two PCA axes explain 42% of the total variability.
NH4incub. is concentration of N-NH4 after incubation; NO3incub. and N_inorg_incub.
have the same meaning for N-NO3 and sum of N-NH4 and N-NO3.
3.2.3. Soil reaction, base saturation and soil carbonates
Soil reaction and degree of base saturation in BL stands varied greatly depending on parent rock. In the Czech Republic, BL grows in extremely acid soils over lying lydite (pH(H2O) = 3.4), eolian sands
(3.8) and Proterozoic schist (3.9), as well as in mildly alkaline soils on
basalt (7.4), limestone (7.4), Ordovician schist (7.5), olivine nephelinite
(7.6), Carbon-Permian sandstone (7.6) and moderately alkaline soil on
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
calcareous sandstone (7.9). However, 69% of BL stands grow in extremely and very strongly acid soils.
Base saturation (BS) also varied greatly—from medium saturation
(31–50%, e.g. Proterozoic schists, eolian sands, lydite or quartzite) to
highly saturated (more than 75%; e.g. granodiorite, spilite, diabase,
amphibolite or calcareous sandstone and limestone). Base saturation
of A-horizon in BL stands was usually very good with an average value
of 74% (Table 2). Low CEC values (less than 12 meq/100 g) were recorded for eolian sands and quarzite sandstone and very high values (more
than 40 meq/100 g) for basic rocks (basalt, spilite, diabase or limestone). Calcium was the most important exchangeable cation with
other elements playing only a marginal role. However, calcium content
varied considerably depending on the type of bedrock (from 264 mg/kg
on eolian sands up to 20956 mg/kg on spilite; Table 2). The highest concentration of Mg2+ was recorded for olivine nephelinite (1945 mg/kg),
and the lowest for eolian sands (13 mg/kg). At nine localities with
either extremely acid or very strongly acid soils on schist bedrock and
clay minerals that served as a source of aluminum, BS was recorded in
the Al cycle. Soils rich in carbonates were described on the following
bedrocks: diabase, spilite, basanite, olivine nephelinite, limestone and
calcareous and Carbon-Permian sandstones. On acid substrates, the carbonates were only present in the upper part of the A-horizon, which
was rich in shells, especially those of the gastropod Helix pomatia.
Expected positive correlations between soil reaction and exchangeable calcium (r.s. = 0.782; p b 0.001), magnesium (r.s. = 0.604
and p b 0.001), CEC (r.s. = 0.547; p = 0.001) or BS (r.s. = 0.933;
p b 0.001), and negative correlations between soil reaction and
exchangeable aluminum (r.s. = − 0.847; p b 0.001) or hydrogen
(r.s. = −0.818; p b 0.001) arise from cation exchange processes (e.g.
Sparks et al., 1996). Soil pH was significantly increasing with the soil
depth (p = 0.01), whereas exchangeable cations showed statistically
insignificant change.
Soil reaction was detected as the most important (p = 0.0001) from
55 studied environmental characteristics to drive the species composition and amount of nitrophytes in BL stands. Other environmental
characteristics were not significant after the influence of pH was filtred
out. Efficiency of constrained axis (pH) was 81% of the total variability
explained by unconstrained axis.
3.2.4. Exchangeable phosphorus
In the stands studied, exchangeable soil phosphorus was present
at a relatively low concentration. The average value of phosphorus
was 20.9 mg/kg (Table 2). The minimum value (under 2 mg/kg)
was recorded in grassy stands on slopes of river valleys. The maximum of 110.9 mg/kg measured in a nitrophilous BL stand dominated
by Anthriscus cerefolium var. longirostris is considerably greater than
other measures.
3.2.5. Total carbon, nitrogen and Corg/N ratio
Total nitrogen content was strongly positively associated with total
carbon (r.s. = 0.983; p b 0.001), because most of both elements come
mainly from organic matter. Carbonates occurred only rarely and in
low concentrations; therefore strongly positive correlation between
total and organic carbon was detected (r.s. = 0.999; p b 0.001). Consistent with the above mentioned findings, maximum values of total C and
N were recorded at the same localities (Table 2)—on eutric Leptosol
with a thick layer of litter from the shrub Grossularia uva-crispa and
dystric Cambisol dominated by Impatiens parviflora (cf. Hofmeister
et al., 2009). High contents of both these elements were recorded in
stands with a low soil reaction, base saturation and mineralization in
which the litter layer was formed by poorly decomposing biomass of
grasses, such as P. nemoralis or Avenella flexuosa. Consistently, both
elements were negatively and significantly associated with soil pH
(Corg: r.s. = −0.599; p b 0.001; N: r.s. = −0.560; p b 0.001). The lowest
values (Table 2) for both total carbon and nitrogen were recorded on
basalt, and in spring also in nitrophilous stands (Fig. 3), probably due
323
to rapid mineralization of organic matter. Both elements significantly
decreased with depth in the soil profile (p b 0.01).
Maximum Corg/N (more than 15; Table 2) was measured in stands
dominated by the grasses Brachypodium sylvaticum, Poa angustifolia
and P. nemoralis growing on green schist and Calamagrostis epigejos
dominated stands on eolian sands (Fig. 2). Minimum values of around
10 were recorded for stands dominated by nitrophytes growing on
basic rocks (Fig. 3) and surprisingly also for dwarf and shrubby stands
(Fig. 1). Corg/N ratio was significantly negatively correlated with pH
(r.s. = −0.615; p b 0.001) and BS (r.s. = −0.561; p = 0.008). Corg/N
ratio usually varied between 10 and 12; higher values were reached
only where the pH(H2O) was lower than 4.3. Low value of Corg/N indicates high quality of soil and rapid rates of decomposition (Killham,
1994; Manzoni et al., 2010). According to our observations, BL produces
relatively small amounts of leaf litter and therefore only a thin layer of
moder or mull-like moder humus develops in stands of this tree.
Composition of the herb layer, especially cover of nitrophilous species
with broad leaves or slope, can be an important factor, as lightweight
BL leaves are drift easily and deposit on lower parts of the slope.
3.2.6. N mineralization, nitrification and ammonification
In the BL stands studied, the rate of N mineralization at the beginning of the growing season differed (Table 2). The maximum rate of
N mineralization (327.1 mg NH4 + NO3–N/kg/28 days) and nitrification (299.6 mg NO3–N/kg/28 days), and a high rate of ammonification (27.5 mg NH4 –N/kg/28 days) was recorded in a stand
growing in moist Eutric Leptosol on spilite with high edaphon activity. Most of the mineralized nitrogen was formed by NO3–N, except
in BL stands at the base of lydite outcrops, where intense mineralization (113.3 mg/kg/28 days) was caused by high ammonification
(108.9 mg/kg/28 days). The intensity of ammonification was negative or near zero except at three localities. This demonstrates that
the majority of the NH4-N input was nitrified during incubation. Immobilization of nitrogen (indicated by negative or very low intensity
of N mineralization, ammonification and nitrification) was recorded
at BL stands with dense grass cover (Arrhenatherum elatius, Bromus
erectus, P. nemoralis, Festuca ovina or A. flexuosa) on totally different
bedrocks (aeolian sand, Proterozoic schist and olivine nefelinite).
From all studied environmental variables including soil properties
(see Chapter 2.4), N mineralization rate, incubated inorganic nitrogen
and nitrates were the best positively correlated with the base saturation
and CEC (Fig. 6). Intensity of N mineralization was significantly positively correlated with the concentration of NO3-N before incubation
(r.s. = 0.453; p = 0.008), and NO3-N (r.s. = 0.688; p b 0.001) and inorganic nitrogen (r.s. = 0.820; p b 0.001) produced during incubation.
Nitrification rate followed closely N mineralization (r.s. = 0.720;
p b 0.001); therefore the quantity of nitrates and inorganic nitrogen
after incubation was also positively correlated with nitrification
(r.s. = 0.800 and 0.677, respectively; both p b 0.001). High initial
concentrations of ammonium were associated significantly with low
ammonification rates (r.s. = −0.709; p b 0.001). Intensity of ammonification decreased significantly with the increase in the rate of nitrification (r.s. = −0.612; p = 0.03). The majority of NH4–N was consumed;
hence these ions were usually not accumulated in the soil profile of BL
stands.
No correlation was detected between number/share of nitrophytes
and nitrification or N mineralization (Fig. 6). Surprisingly, high rates of
both characteristics were detected in nitrophilous as well as dwarf
and shrubby stands on granodiorite outcrops. Unlike laboratory conditions (i.e. sufficient soil moisture—60% of maximum capillary capacity,
and high temperature—28 °C), in natural conditions low soil moisture
(8%) probably prevented the development of nitrophilous species.
3.2.7. Inorganic forms of nitrogen
Concentrations of all forms of nitrogen decreased significantly with
the soil depth (p b 0.01). The volume of nitrates reached very different
324
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
values, especially in September 1999, when samples were collected
after a long dry period. Nitrate concentration was significantly positively correlated with CEC (r.s. = 0.569; p = 0.004) and soil moisture
(r.s. = 0.635; p b 0.001). We detected no nitrates in eolian sands containing little moisture with low N mineralization rate, and maximum
NO3–N (203.4 mg/kg; Table 2) in spilite debris with the highest soil
moisture content and N mineralization rate. Between September and
April, the concentration decreased at most localities (from 24.3 mg/kg
to 12.1 mg/kg on average; Table 2), probably due to intake of spring
species of plants and leaching from the A horizon. At most localities,
the contribution of nitrates to the total nitrogen was higher than that
of ammonium ions, especially in BL stands in which the herb and
shrub layers were dominated by nitrophytes (Figs. 3 and 6). High
number and ratio of nitrophytes were typical for nitrophilous BL forests
on alkaline soils (Fig. 6).
The ammonium concentrations varied less compared to nitrates and
were markedly low in most BL stands. High values were usually associated with the clay mineral content in bedrock. The quantity of NH4–N
increased significantly from September 1999 to April 2000 (6.8 to
7.8 mg/kg on average; minimum in autumn on granodiorite rock
steppe—1.5 mg/kg; maximum in spring on Proterozoic schists
45.2 mg/kg; Table 2), probably due to decomposition and mineralization of dead organic matter and weathering of bedrock caused by
large fluctuations in temperature. Concentration of ammonium in the
A-horizon was correlated significantly positively with soil moisture
(r.s. = 0.597; p b 0.001), total nitrogen (r.s. = 0.667; p b 0.001) and
both total (r.s. = 0.703; p b 0.001) and organic (r.s. = 0.692;
p b 0.001) carbon, and negatively with soil reaction (r.s. = − 0.458;
p = 0.007). The relationships described above are confirmed by
Fig. 6, from which it is evident that NH4–N was accumulated in skeletal acid soils rich in organic matter with a high content of organic
carbon and total nitrogen. Such soils are preferred by open and
mesic BL stands where poorly decomposing biomass of grasses
A. flexuosa and P. nemoralis dominate and nitrophilous species are
rare (see Fig. 6).
3.3. Comparison of BL soil types from different parts of its distribution range
The variability of soil types in BL stands corresponds to the diversity
of habitats which BL is able to invade. Tree vegetation and soils in BL
stands in the northern Appalachian Mountains (part of its native
range) and in its secondary range in Central Europe are very similar
(Hagner, 1999; IUSS, 2006). The most common soil type in both is
Cambisol with incipient soil formation. Cambisols dominate both in
temperate and boreal regions but are less common in the tropics and
subtropics (IUSS, 2006). Outside Europe, BL is common in temperate
Asia, Australia, New Zealand, northern and southern Africa and temperate South America (Weber, 2003), but there is no information on the
soils on which they grow in these areas. In the sub-tropical zone in
China, BL occurs on Cambisols (Yanna et al., 2013).
The humid climate in the southern Appalachians (North Carolina;
Boring and Swank, 1984a; Montagnini et al., 1989; Huntley, 1990)
differs from the conditions prevailing in the Czech Republic. In the
southern Appalachians BL stands dominate on Luvisols and Acrisols
whereas in the Czech Republic with moderate climate, these are rare
(Haplic Luvisols cover 7% and Albeluvisols 2%; Fig. 5). In Central Spain
with a Mediterranean climate, BL occurs mainly in riparian forests and
forest plantations on Calcic Luvisols and Fluvisols (Calcaric and Eutric;
Castro-Díez et al., 2009). In north-eastern Slovenia, Ivajnšič et al.
(2012) report that the most frequently invaded soils are Stagnosols
(38%), followed by Fluvisols (21%) and Eutric Cambisols (17%). In the
Czech Republic, Fluvisols (7% of the localities studied; Fig. 5) occur in
BL stands growing at the bases of slopes, mostly in river and stream valleys. Both in its native range (Huntley, 1990) and the Czech Republic, BL
rarely grows on poorly drained, compact plastic or waterlogged soils.
The reason could be that nitrogen fixation is inhibited in such
environments. Aleksandrova et al. (1994) report that nodule formation,
nitrogenase activity and ureide transport in BL organs are inhibited and
the total biomass per plant is lower when growing in alluvial-meadows
than in sandy soils.
In continental Europe, BL stands usually thrive on Arenosols developing in sand dunes (Keresztesi, 1988; Šimonovič et al., 2001; Vrbek
and Pilaš, 2011) or on river terraces formed from loamy sand or coarser
fragments of up to 35 percent by volume, as in the Elbe lowlands (Czech
Republic, our study; Fig. 2). In accordance with Samarakoon et al.
(2013) we found that BL is able to tolerate the temporary filling of
pores in sandy soil with water and is to some degree resistant to scouring (i.e. removal of substrate from the root-anchoring zone and the
exposure of its roots).
Although Fowells (1965) and Huntley (1990) state that very dry
sites and soils less than 24 in deep (aprox. 60 cm) are unsuitable for
BL in its native range, in the Czech Republic it often invades such habitats (Fig. 1). Leptosols were present in 35% of the stands studied
(Fig. 5), of which the most frequent were lithic and typic Leptosols
with a dry and narrow A horizon. BL also occasionally grows in rocky
soils (foothill of the Southern Alps; Cierjacks et al., 2013) and in the Vesuvius National Park (Southern Italy) it occurs on shallow Andosols with
weak profile differentiation (De Marco et al., 2013). To conclude, in its
secondary range BL often grows in soils that are drier and shallower
than those in its native range (Huntley, 1990).
3.4. Comparison of BL physical–chemical soil properties in different parts of
its distribution range
Even though BL can markedly change the soil environment, the
physical-chemical properties of soils in areas where BL occurs have
never been discussed comprehensively. Some authors deal with particular aspects, such as nitrogen fixation (e.g. Boring and Swank, 1984b;
Danso et al., 1995; Noh et al., 2010), soil nitrogen and its transformation
(e.g. Guofan and Tingxiu, 1991; Moon, 1999; Rice et al., 2004; Malcolm
et al., 2008; Castro-Díez et al., 2009), carbon dynamics (e.g. Harris and
Riha, 1991), litter decomposition (e.g. White et al., 1988; Harris and
Safford, 1996; Yanna et al., 2013) and microbial analysis (e.g. Eaton
and Farrell, 2004; Landgraf et al., 2005; Wang et al., 2012). There are
several comparisons of BL invaded and non-invaded sites (e.g.
Montagnini et al., 1986, 1991; Dzwonko and Loster, 1997; Rice et al.,
2004; Landgraf et al., 2005; Von Holle et al., 2006; Malcolm et al.,
2008; Castro-Díez et al., 2012; De Marco et al., 2013; González-Muñoz
et al., 2013; Zhou et al., 2013; Cools et al., 2014) or BL stands and native
grasslands (Qiu et al., 2010).
Soil characteristics of BL stands in its native range were investigated
in detail only in its southern part (western North Carolina, the Coweeta
LTER, 2 sites) with humid continental climate (e.g. Boring and Swank,
1984a,b; Montagnini et al., 1986, 1989, 1991; White et al., 1988) and
are missing from the northern part of native range that would be comparable to the Central Europe regarding the climate, tree vegetation and
soil types. The data available from the secondary range also do not cover
the whole variability of BL soil conditions and describe situation at
isolated sites only. Additionally, different analytical methods were
used between the current and previous studies, further preventing
such statistical analysis. This makes the statistical comparison of our results to published data and comparison of primary and secondary range
soil conditions problematic. The biggest barrier is the application of various extraction agents, simple and composite, with diverse chemical
composition (neutral electrolytes, chelate solutions, acids and their
salts, oxidation reagents), which show different extraction impact (cf.
Sparks et al., 1996; Ure, 1996; Zbíral, 1999; Kulhánek et al., 2009;
Zbíral and Němec, 2009). Moreover, the nutrient availability in forest
floor varies depending on many factors such as climatic conditions,
stand age or vegetation season (e.g. Nadelhoffer et al., 1984; Pastor
and Post, 1986; Brais et al., 1995). Considering the above mentioned,
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
our results could be compared with published data from primary and
secondary range on a qualitative basis only.
3.4.1. Soil moisture
Variability in soil moisture influences mineralization of organic matter and release of mineral nitrogen into the soil solution (e.g. Cassman
and Munns, 1980; Stark and Firestone, 1995), as well as nutrient
leaching due to the weathering of the bedrock. Under BL stands, the
soil can quickly dry out because of the later and less dense foliation;
however a large root system enables BL to obtain water, even in dry
stands (e.g. Göhre, 1952). Větvička (1965) found a greater uptake of
water by roots in the upper soil layer (5 cm) in BL compared to both
oak and mixed BL-oak forests.
3.4.2. Soil reaction and base saturation
Black locust is able to grow in a variety of soil conditions (Table 2). In
its native range, soil reaction ranges from extremely acid (4.0) to moderately alkaline (8.2; Fowells, 1965; Vogel, 1981) with no apparent
effect on growth (Huntley, 1990), whereas in its secondary range it is
3.2 (extremely acid soil reported by Kowarik, 1992 for Germany) to
7.9 in Europe (moderately alkaline by Castro-Díez et al., 2009 for Central
Spain and the Czech Republic) and 8.8 in the world (strongly alkaline
soils in China according to Yanna et al., 2013; Table 2). Using fertilizer
experiments, Fowells (1965) observed decrease in the growth of BL in
pH above 6.9, however in the Czech Republic BL stands grew well in
soils ranging from 3.4 to 7.9. Medium saturated soils (Table 2;
Montagnini et al., 1986) were recorded for Coweeta LTER (BL in its
native range), while in the Czech Republic base saturation reached up
to 100%, i.e. highly saturated alkaline soils.
3.4.3. Phosphorus and nitrogen
Forest soils are usually poor in terms of NH4–N and NO3–N; symbiotic N2 fixation by BL increases the amount of available nitrogen for plants
(Boring and Swank, 1984b). Deficiency of phosphorus in BL stands
decelerates and finally inhibits symbiotic fixation of atmospheric nitrogen (Liu and Deng, 1991), even before causing tree growth to cease
(Úlehlová, 1989). On basic soils, phosphorus can be a limiting element.
The low measured concentration of phosphorus in our research could
be a consequence of a low amount of apatite in the bedrock, higher
leaching of PO4-P ions (cf. Van Miegroet and Cole, 1984; Montagnini
et al., 1989), or its intensive uptake by BL roots and accumulation in
leaves (cf. Montagnini et al., 1989). In the case of a shortage of phosphorus high nitrogen availability does not lead to higher vitality of an
ecosystem (Van Oorschot et al., 1997).
The content of NO3–N was higher in BL soils than in the native
forests at Coweeta LTER (BL native range), whereas there were no
differences in the NH4-N concentrations, total nitrogen and C/N in
these soils (Montagnini et al., 1989). Nitrate concentrations and nitrification potentials in BL stands growing on slopes of various aspects were
similar (Montagnini et al., 1986). The values cited from the native range
by Montagnini et al. (1986) –4.45 mg/kg in March and 3.25 mg/kg in
July were much lower compare to our measured average NO3–N concentrations (12.13 mg/kg in April and 24.34 mg/kg in September),
and Dzwonko and Loster (1997) values from Poland (35.5 mg/kg in
September; Table 2). Nevertheless, comparison with other studies is
difficult because the concentration of soil NO3–N depends on many factors such as climatic conditions and stand age, and varies greatly both
spatially and temporally even at the same locality (e.g. Boring and
Swank, 1984a; Nadelhoffer et al., 1984; Cambardella et al., 1994;
Montagnini et al., 1991; Brais et al., 1995; our unpublished data).
Montagnini et al. (1991) state that the seasonal pattern is caused by
interactions between nitrification, plant and microbial uptake, denitrification and leaching, with the lowest nitrate concentrations reached
during winter due to limited nitrifying activity, and in July due to greater
plant uptake. Also dry conditions minimize the loss of nitrates through
leaching and reduce the rate of mineralization of organic matter (e.g.
325
Montagnini and Sancho, 1994). Peaks in October and November are
caused by high production of NO3–N and decreased uptake by plants
(Montagnini et al., 1991). According to these findings, our measurements were higher in September (end of the growing season) than in
April (beginning of the growing season, following a long and cold winter). Montagnini et al. (1989) in accordance with our research recorded
decrease of NO3–N, NH4–N, total nitrogen and carbon with soil depth.
3.4.4. Nitrification and its acidifying effect
Consistent with Montagnini et al. (1986), net N mineralization was
very closely correlated with nitrification rate. In both the native range
and our study area, all the NH4–N was consumed during incubation
and the rate of ammonification decreased significantly with increasing
nitrification. Montagnini et al. (1989) conclude that the availability of
ammonium in BL soils is the main factor controlling nitrification, ammonification is more important than exchangeable Ca2 + or pH
(Montagnini et al., 1986). In comparable periods, the value of total nitrogen detected in the native range (Boring and Swank, 1984a;
Montagnini et al., 1986) reached the minimum measured in our study
(Table 2). Equally, the nitrification and mineralization rates were near
the lower limit (Table 2), which may be due to the different ages of
the BL stands—12 years in Coweeta (Montagnini et al., 1986) and
more than 40 years in our study. Consistent with Castro-Díez et al.
(2009), immobilization of nitrogen was detected in dense grassy BL
stands with a large input of carbon-rich plant material. Harris and
Riha (1991) report an inverse relationship between C and N mineralization, which eventually resulted in N immobilization.
Van Miegroet and Cole (1984) showed that acidification of the top
10 cm of soil under red alder (Alnus rubra, a nitrogen fixing tree) coincided with intense nitrification, which provided a major source of H+
from the dissociation of the strong HNO3 acid produced by this microbial oxidation reaction. In the BL stands Montagnini et al. (1986) detected
the highest nitrification activity in the top 15 cm of the soil. Nitrification
was found to be a greater source of acidification than deposition of
atmospheric H+, even in areas markedly affected by acid precipitation
(Van Miegroet and Cole, 1984). Simultaneously, NO−
3 leaching could
result in an increase in the concentration of N in the groundwater. For
the foothills of the Cascade Mts. (near Seattle, WA), Van Miegroet and
Cole (1984) reported that the presence of large amounts of mobile
NO−
3 in solution triggered accelerated cation leaching and caused selective redistribution of exchangeable Ca2+ from A to B horizon. While this
acidifying effect of nitrification was not detected in the BL stands in its
native range (Montagnini et al., 1991), the soil reaction in the Czech
Republic (this study) significantly increased with the depth of the soil
profile, whereas redistribution of exchangeable cations was not recorded. In forests composed of nitrogen non-fixing tree species, Augusto
et al. (2002) mentioned other possible sources of acidification in topsoil
as litter decomposition, deposition and root exudates. The tree layer
species composition is very important, because leaf litterfall rich in
Ca2+ and Mg2 + (e.g. ash, lime or maple) can reduce acidifying effect
and increase the nutrient availability in topsoils (Guckland et al.,
2009; Langenbruch et al., 2012). Although BL leaves have similarly
high cation content (Montagnini et al., 1989; Rice et al., 2004), in our
study there were no signs of acidifying effect being reduced in the topsoil. The cause of a BL acidifying effect has not been explained yet.
4. Conclusions
In its secondary range, BL grows in a wide range of soils. The most
common soil types are shallow young soils, such as Cambisols,
Leptosols, Arenosols and Fluvisols with incipient soil formation. Thriving BL stands were also recorded growing in deep and rich Chernozems
(in continental Europe), and Acrisols and Luvisols with an Argic illuviation horizon (in sub-humid and wet subtropical climates). The best
conditions for the growth of BL appear to be sandy soils like Arenosols.
326
M. Vítková et al. / Science of the Total Environment 505 (2015) 315–328
BL is also able to tolerate diverse soil chemistry. Soil reaction varied
from extremely acid (pH(H2O) 3.2 in the secondary and 4.0 in the native
range) to moderately and strongly alkaline (8.2 in the native and 8.8 in
the secondary range). Also the base saturation varied greatly—from
medium (30–50%) to highly saturated soils up to 100%. Although
Fowells (1965) mentions that limestone soils are the best for BL growth,
in the Czech Republic BL produces similar high-quality stands on
basaltoids and aeolian sands. Water supply and soil aeration seem to
limit colonization by BL more than the soil chemistry. In both its native
and secondary range, BL prefers well aerated and drained soils. It can
survive the drying out of the soil profile and avoids compact soils and
those that are waterlogged for long periods of time, probably because
this inhibits nitrogen fixation. Low soil moisture on slopes has considerable effect on the vigor of BL and causes scrubbiness and a decline in
competitiveness in favor of drought-adapted native shrubs such as
Crataegus sp., P. spinosa and Rosa sp.
High flexibility of BL makes it favorable species for forest planting;
however it poses serious problems in nature conservation, lowering
biodiversity, spreading spontaneously into the surroundings, and
preventing the natural ecosystem restoration.
Acknowledgments
The authors would like to thank J. Hofmeister for help with the
statistical evaluation of the data set, M. Albrechtová for advice on the analytical methods used in the different studies, Tony Dixon and Grant
Hamilton for editing the English, and anonymous reviewers for valuable
comments on the manuscript. This study was supported by the longterm research development project no. RVO 67985939 and by the
Praemium Academiae award to P. Pyšek from the Academy of Sciences
of the Czech Republic.
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