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Available at www.sciencedirect.com
http://www.elsevier.com/locate/biombioe
Potential benefits of commercial willow Short Rotation
Coppice (SRC) for farm-scale plant and invertebrate
communities in the agri-environment
Rebecca Rowe 1, Mick Hanley 3, Dave Goulson 4, Donna Clarke 1,
C. Patrick Doncaster 2, Gail Taylor*
University of Southampton, Faculty of Natural and Environmental Sciences, Life Sciences Building, Southampton, S017 1BJ, United Kingdom
article info
abstract
Article history:
The cultivation of bioenergy crops (BECs) represents a significant land-use change in agri-
Received 20 April 2009
environments, but their deployment has raised important issues globally regarding
Received in revised form
possible impacts on biodiversity. Few studies however, have systematically examined the
12 August 2010
effect of commercial scale bioenergy plantations on biodiversity in agri-ecosystems. In this
Accepted 18 August 2010
study we investigate how the abundance and diversity of two key components of farmland
Available online xxx
biodiversity (ground flora and winged invertebrates) varied between mature willow Short
Rotation Coppice (SRC) and two alternative land-use options (arable crops and set-aside
Keywords:
land). Although the abundance of winged invertebrates was similar across all land-uses,
Bioenergy
taxonomic composition varied markedly. Hymenoptera and large Hemiptera (>5 mm) were
Biodiversity
more abundant in willow SRC than in arable or set-aside. Similarly although plant species
Willow SRC
richness was greater in set-aside, our data show that willow SRC supports a different plant
Land management
community to the other land-uses, being dominated by competitive perennial species such
Set-aside
as Elytrigia repens and Urtica dioica. Our results suggest that under current management
Semi-natural habitat
practices a mixed farming system incorporating willow SRC can benefit native farm-scale
biodiversity. In particular the reduced disturbance in willow SRC allows the persistence of
perennial plant species, potentially providing a stable refuge and food sources for invertebrates. In addition, increased Hymenoptera abundance in willow SRC could potentially
have concomitant effects on ecosystem processes, as many members of this Order are
important pollinators of crop plants or otherwise fulfil an important beneficial role as
predators or parasites of crop pests.
ª 2010 Published by Elsevier Ltd.
* Corresponding author. Tel.: þ44 (0) 2380592335.
E-mail addresses: [email protected] (R. Rowe), [email protected] (D. Clarke), [email protected] (C.P. Doncaster), [email protected].
uk (G. Taylor).
1
Tel.: þ44 (0) 23 8059 8429.
2
Tel.: þ44 (0) 2380594352.
3
Present address. University of Plymouth, School of Biological Sciences, Drake Circus, Plymouth, PL4 8AA, United Kingdom.
Tel.: þ44 (0) 1752 238319. [email protected]
4
Present address. University of Stirling, School of Biological & Environmental Sciences, Stirling, FK9 4LA, United Kingdom.
Tel.: þ1786 467759. [email protected]
0961-9534/$ e see front matter ª 2010 Published by Elsevier Ltd.
doi:10.1016/j.biombioe.2010.08.046
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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1.
b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
Introduction
Increased use of organic farming practices, genetically modified crops and the implementation of agri-environment and
other biodiversity enhancement schemes have brought about
wide-scale changes to farming throughout the developed
world [1,2]. More recently, the cultivation of dedicated nonfood bioenergy crops (BECs) for power generation and biofuel
production has raised concerns about potential effects on
biodiversity in the agricultural environment [3e5]. Current
emphasis centres on biodiversity loss in developing nations,
but there have been significant shifts towards the cultivation
of BECs in Europe [6], North America [7], and Australasia [8].
Although a number of plant species are utilized as BECs, it is
the use of perennial grasses and fast growing woody crops e
the so called “second generation crops” that pose the greatest
changes in farm practices and have the largest potential to
impact on biodiversity in the agri-environment [4,9].
Willow (Salix spp.) Short Rotation Coppice (SRC) is one of
the most widely planted BECs in Europe [6,10]. It has been
cultivated in the UK since the late 1980s, but the area of land
dedicated to willow SRC has increased dramatically in recent
years from under 1000 ha in 1999 to over 5000 ha in 2007 based
on planting grant applications [11]. Long-term predictions
suggest that 27 000e70 000 M ha of woody crops could be
required by 2050 to meet bioenergy commitments, representing 11e29% of land cover in the UK [4].
Most research to-date suggests that SRC willow has positive
effects on biodiversity [4]. However, these studies often focus
on charismatic groups of species such as song birds [12,13] and
butterflies [9,14], or potential pest species such as canopy invertebrates [15,14]. Moreover, few studies have examined how SRC
affects species composition or abundance in comparison to the
common alternative forms of land-use in the agri-environment
(see [14]). This prevents any realistic assessment of the biodiversity implications of SRC expansion in Europe and beyond.
A further problem associated with many earlier studies on
biodiversity within SRC plantations is that study sites were
often located within small, non-commercial research-scale
plantations, under 3 ha in area and managed in a way
inconsistent with commercial SRC plantations (e.g. different
harvesting cycles, greater mix of willow cultivars/clones per
field, and greater range of age classes). Cunningham et al. [14]
was one of the few studies to address this issue by selecting
only large commercial sites. However, all sites were newly
established (maximum plantation age of 4 years), and therefore, failed to reflect the true nature of mature willow SRC
fields that may remain in use for up to 25 years [16].
The aim of this study was to compare biodiversity impacts
of mature, commercial, large-scale willow SRC plantations
with that observed in the two main alternative land-use
options in the UK, arable and set-aside. Set-aside (land taken
out of food production) was, until 2008, required under EU
Common Agricultural Policy (CAP) in order to regulate food
production. However, under the provisions of the CAP, setaside could be used for the production of BECs, and thus it was
particularly susceptible for conversion [17]. Currently, setaside requirement is set at zero percent in the EU [18].
Consequently, BECs may now be an attractive option for any
low grade farmland that, in the past, land owners often used
to meet set-aside requirement.
Vascular plant abundance and diversity were investigated
as they represent a significant biodiversity component within
the agri-environment [19]. In addition, vascular plants provide
shelter and food for a range of other species, making them
critical to species diversity at the community level [20]. We
also assigned plant species to life-history groupings based on
[21] C-S-R strategy scheme, to make the results of this study
comparable across geographic regions and provide an insight
into the ecological processes affecting plant community
development [21,22]. We assessed the abundance and diversity of winged invertebrates since this group of organisms has
received remarkably little attention in previous studies of SRC
yet comprises the bulk of terrestrial biodiversity and provides
crucial ecosystem services as pollinators and predators of
farm pests [23].
2.
Methods
Field sites were selected primarily on criteria designed to
ensure that sites represented mature commercial plantations.
These criteria included:
Commercial plantations managed in accordance with
current Department for Environment, Food and Rural
Affairs (DEFRA) guidelines [16]
Individual fields greater than 5 ha in size
Sites at least 5 years old, which had completed at least one
harvest cycle
Control fields of arable land and set-aside available close to
plantations
Plantations and control fields to be uniform in shape (i.e.
standard field layout rather than narrow strips or convoluted in shape).
In total, three sites were selected in north Nottinghamshire, UK. SRC plantations ranged from 5 ha to 9 ha, and were
established between 1998 and 2000 (Table 1). In all cases
plantations consisted of a mix of 5 varieties of willow, containing approximately 30% Tora, with the remaining mix
consisting of equal selection of three varieties from Ulv, Olof,
Jorunn, Jorr, and a small amount <10% of Bowles Hybrid.
Arable and set-aside fields were selected for their proximity
and similarity (size and shape) to the SRC plantations. Arable
fields which had been cultivated for cereal crops were
selected, as cereals represent the highest proportion of arable
land-use in Great Britain [24]. In all cases the arable fields had
been cultivated with barley and had been harvested, between
one and two weeks previously. The fields had however yet to
be ploughed being stubble at the time of the study (August
2006) thus the ground flora was relatively undisturbed. The
selected sites were relatively uniform in shape and all were
previously arable land (Table 1).
2.1.
Invertebrate diversity and abundance
Winged invertebrates were sampled using double-sided
yellow sticky traps 22 cm 41 cm (BC28211, Agrisense-BCS
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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Table 1 e Field site details giving grid references, field size, establishment year, (for willow year of planting for set-aside
first year of registration) and date of last harvest. All sites were located in north Nottinghamshire and were selected based
on criteria relating to age, and size of plantation, and location of plantation in relation to control fields. In all cases previous
land-use was arable.
Site
Land-use (plots)
Location (WG84 DMS)
North
1
2
3
Willow SRC
Arable (barley)
Set-aside
Willow SRC
Arable (barley)
Set-aside
Willow SRC
Arable (barley)
Set-aside
53:21:23.18
53:20:42.22
53:19:44.93
53:25:59.40
53:26:02.47
53:26:16.19
53:26:26.09
53:26:28.25
53:26:22.62
Size (ha)
Year established
Date of last harvest
7.67
20.01
3.82
9.00
5.32
6.69
5.75
10.00
5.87
2000
N/A
2004
1998
N/A
2004
1998
N/A
2001
2005
July 2006
N/A
2004
July 2006
N/A
2004
July 2006
N/A
West
0:59:57.60
0:59:42.28
0:58:56.42
0:48:6.62
0:48:07.54
0:46:58.62
0:47:20.44
0:46:57:12
0:47:03.05
Ltd., Treforest Industrial Estate, Pontypridd, Mid-Glamorgan,
UK). As described previously [25,26]. Yellow traps were
selected over other colour options as they are considered to be
effective over the widest range of invertebrate species [27]. To
ensure samples were taken from an area as wide as possible,
fields were divided into equal quarters. One side of the field
was randomly selected and two transects along the center of
the quarters (running from headland into to the crop at right
angles crop edge). The third transect was then placed on the
opposite side of the plantation at the intersection of the two
remaining quarters. Sampling points were located along each
transect in the headland, 5 m, 25 m, 50 m and 100 m into the
cultivated area, apart from site 3 where the centre of the SRC
was at 61 m, a central sampling point was used, both in the
plantation and in the paired arable and set-aside fields.
As height has been reported to affect sticky trap efficiency
[25] a set of three traps were installed at each sampling point,
0.10 m, 1 m and 2 m above ground level. This ensured that at
least one trap in each land-use type was close to the vegetation canopy. Each set of three traps was suspended between
two bamboo canes such that the 22 cm edge of each trap was
parallel to the ground. To prevent vegetation adhering to the
traps and thereby reducing their efficiency, each trap was
surrounded by an open-ended tube made from galvanised
wire netting (mesh size of 50 mm, Gardman, Moulton,
Spalding, Lincolnshire, UK).
Traps were installed in each site over a 3-day period in
August 2006, with each land-use (willow, arable, or set-aside)
taking a full day to set up. Each trap was left in place for 144 h,
before being collected, wrapped in cling film and frozen at
20 C. All invertebrates over 5 mm in length were identified
to Order. For invertebrates less than 5 mm in size, each side of
the trap was divided into a 2 2.1 cm grid and all individuals
within 10 randomly selected squares per side (5% of the total
trap area) were identified to Order using a dissection microscope. Thus results for some Orders were divided into two size
classes; referred to as ‘large’, (over 5 mm) and ‘small’ (under
5 mm). All individuals present on a given trap (regardless of
size) were counted to give a total winged invertebrate abundance per trap.
Statistical analyses were performed in Minitab version 15
after normalising residuals with a square root transformation.
The effects of land-use, distance into the crop, and trap height
on the abundance of winged invertebrates were examined
using the following split-plot nested ANOVA model (henceforth referred to as model 1):
Abundance ¼ H3 jD5 jT03 F01 B03 L3
where prime identifies a random factor, subscript refers to
number of factor levels, “j” to “cross-factored with”, and “(“ to
“nested in” [28]. H ¼ height, D ¼ distance into crop (headland,
5 m, 25 m, 50 m, and 100/61 m), T0 ¼ transect, F0 ¼ field,
B0 ¼ blocking factor site, and L ¼ land-use. With a single field
for each of the nine B0 * L combinations, fixed main effects and
their interactions were each tested against their respective
interactions with site (which were not themselves testable
because fields were not replicated for each land-use within
the three site blocks). Although the low site replication gave
few error d.f. for testing the land-use main effect, power to
detect an effect was improved indirectly by the error variation
being estimated from replicate transects. Larger numbers of
error d.f. were available for testing land-use interactions with
other treatment factors.
2.2.
Ground flora
To account for the planting pattern in Willow SRC plantations
a 2 m 2 m quadrat was used to allow sampling of both
a section of the tramlines (1.5 m gap between double rows of
willow stools, used for machinery access) and intra-stool area
[29]. Within each quadrat, the cover of each component species
was recorded based on the Domin scale, excluding Bryophytes
[30]. Floral surveys were conducted during August 2006.
Fields were divided into equal quarters and transects positioned in the center of each quarter (giving four transects) Within
each transect, sample points were the same as the winged
invertebrates but with an additional sampling point included
at the edge of the cultivated area. The number of quadrats were
set to allow 80 m2 of cultivated area to be surveyed, an area
equivalent to that recommended for surveying the herb layer in
National Vegetation Classification (NVC) surveys and similar
to that used in previous studies [14,15].
A sample of above ground plant biomass was also taken
from three (randomly selected) ground flora transects. For
each sample 0.25 m2 of above ground biomass was collected
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
from each quadrat, dried at 80 C for 48 h (until no additional
weight loss was seen) and weighed. Plant species recorded
within each quadrat were designated attributes for three plant
strategies: life history (annual or perennial), life form (grass or
forb), and establishment strategy (C-S-R) [21]. Establishment
strategies were then further grouped into four groups, C
competitive species, CSR generalised species, S stress tolerant
species, R ruderal species (following [22]). Prior to analysis of
ground flora diversity, plant strategies, and dominant species,
Domin cover values were transformed into percentages using
the protocol described by Godefroid et al. [31].
Following square root transformation to ensure homogeneity of variances, the effects of land-use and distance into
the crop on plant species richness, diversity and biomass were
examined using the following split-plot nested ANOVA
(henceforth referred to as model 2):
Richness ¼ D6 jT04 F01 B03 L3
Diversity ¼ D6 jT04 F01 B03 L3
Biomass ¼ D6 jT03 F01 B03 L3
As for model 1, fixed main effects and their interactions
were each tested against their respective interactions with the
random variable site.
Due to variation in the total cover between land-uses,
direct comparisons between plant strategies based directly on
percentage cover were inappropriate. Therefore, the level of
cover for a given plant strategy (Si) was calculated as a fraction
of total cover within each quadrat (equation (1)) [22].
Si ¼ Ai =T
(1)
where Ai is the total cover per quadrat of a given strategy
division (e.g. annual, perennial, e.t.c), and T is the total floral
cover per quadrat.
To improve normality of residuals, the fraction of cover at
each sampling location (i.e. headland, 0 m, 5 m, 25 m, 50 m,
and 100/61 m) was averaged across all four transect per field
given mean value per distance. Means were then arcsine
transformed prior to analysis. Due to limited floral cover in
arable land a limited number (maximum of three) sampling
location had no cover. In these cases values were replaced
with average values from the remaining two sites of same
land-use. All strategies with the exception of Sþ which, due to
rarity was not suitable for statistical analysis, were examined
using the following split-plot nested ANOVA (henceforth
referred to as model 3).
Strategy ¼ D6 jF01 B03 L3
Data manipulation was conducted in MS-Excel 2007 and
statistical analysis in Min-tab 15.
3.
Results
3.1.
Winged invertebrates
The abundance of winged invertebrates was significantly
influenced by both trap height and distance into the crop
within the different land-use types (Tables 2e4), In contrast to
arable and set-aside land, in which abundance decreased with
height, invertebrate abundance in willow SRC increased from
0.10 m to 1 m, and remained high at 2 m (Table 4). Invertebrate
abundance within willow SRC headlands was also higher than
in the other land-uses, and higher than in the crop area of the
willow SRC (confirmed by removal of willow data F4,8 ¼ 0.15,
P ¼ 0.960, and headlands data F3,6 ¼ 0.72, P ¼ 0.577, Table 3).
Invertebrate abundance in the other land-uses however, was
not affected by distance into the crop as confirmed by the
removal of the willow data (LcD interaction: F4,8 ¼ 1.43,
P ¼ 0.31).
3.2.
Distribution of winged invertebrate Orders
Fourteen arthropod Orders were observed across all sites,
however, statistical analysis was only applied to the seven
most abundant Orders (Table 2). The remaining Orders were
excluded due to low sample sizes. The abundance of large
Hymenoptera, small Hymenoptera and large Hemiptera were
higher in willow SRC than in the alternative land-uses
(Tables 2 and 3). The remaining Orders showed similar
abundance in all land-uses (Table 2). In many cases however,
land-use had a significant effect as part of an interaction with
height and/or distance. For example, small Diptera (<5 mm)
and large Coleoptera, although common in the headlands of
SRC, were much less abundant in the SRC crop than within
the other land-uses (Table 4). Thysanoptera were also more
abundant in SRC headlands, (Table 4) but their abundance
within the crop remained similar between the land-uses even
with the exclusion of the headland data (F2,4 ¼ 3.37,
P ¼ 0.139).
Height and land-use interactions were also apparent for
Hymenoptera, small Diptera, and Lepidoptera (Tables 2 and 4).
For the most part, the effects of height on these Orders were
largely in accord with the effect on total winged invertebrate
abundance (Table 3). Lepidoptera however, showed a markedly different pattern, with a single peak in abundance at
0.10 m in set-aside, compared to a uniformly low abundance
at all other locations (Table 3). Large Diptera and small Hemiptera were affected only by height (Tables 2 and 3), with
similar overall abundance in each land-use type.
3.3.
Ground flora species richness, biomass and
diversity
Interactions between land-use and distance were present for
species richness, ground flora biomass and diversity (Table 5).
Post hoc testing showed species richness, biomass and diversity to be similar in the headlands of all three land-uses (“L”
effect for Species Richness: F2,4 ¼ 1.42, P ¼ 0.342; biomass:
F2,4 ¼ 1.11, P ¼ 0.415; diversity: F2,4 ¼ 2.87 P ¼ 0.169). Within the
cultivated area (25 m), however, species richness was highest in set-aside land followed by willow SRC and finally arable
land (L effect: F2,4 ¼ 17.45, P ¼ 0.011, Fig. 1A). At all distances
ground flora biomass was similar in willow SRC and set-aside
(Table 5, Fig. 1B), but much reduced in the cultivated area of
arable land (Fig. 1B). Within the cultivated area the Shannon
diversity index was highest in set-aside land, with willow SRC
and arable land showing surprisingly similar levels of
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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573
574
575
576
577
578
579
580
581
582
583
584
585
L
D
D*L
H
H*L
H*D
H*D*L
Factor
F
P
MS
F
P
MS
F
P
MS
F
P
MS
F
P
MS
F
P
20.97
125.54
215.11
1662.5
647.44
36.58
41.74
0.04
3.49
2.93
28.22
10.37
1.19
1.81
0.958
0.062
0.032*
0.004*
0.003*
0.365
0.075
473.92
7.02
7.02
65.21
21.61
1.24
3.56
4.82
1.52
0.73
47.20
1.82
0.43
1.21
0.086
0.285
0.661
0.002*
0.218
0.884
0.314
86.49
6.87
20.55
68.38
25.74
2.58
2.21
4.18
4.88
7.30
11.13
5.36
1.48
2.29
0.105
0.027*
0.001*
0.023*
0.021*
0.239
0.023*
64.92
4.00
0.85
3.94
3.43
0.36
0.42
13.27
11.70
0.95
21.99
6.91
0.57
1.51
0.017*
0.002*
0.506
0.007*
0.010*
0.785
0.156
198.63
3.01
1.59
7.29
5.61
1.52
0.68
16.25
1.24
1.00
5.41
4.30
3.02
0.96
0.012*
0.366
0.471
0.073
0.038*
0.028*
0.515
42.17
0.54
1.38
5.06
2.14
0.15
0.61
14.94
0.63
2.12
18.90
2.02
0.26
1.21
0.014*
0.653
0.095
0.009*
0.184
0.969
0.310
Hemiptera <5 mm
DF
2,4
4,8
8,16
2,4
4,8
8,16
16,32
Coleoptera >5 mm
MS
F
P
11.00
0.50
0.64
7.26
1.19
0.66
0.35
0.84
0.93
0.46
53.07
1.64
1.60
0.92
0.494
0.495
0.866
0.001*
0.255
0.202
0.557
MS
F
32.21
2.50
1.62
1.65
2.50
0.37
0.39
4.57
3.59
4.88
0.98
2.70
1.14
0.87
Lepidoptera >5 mm
Thysanoptera
P
0.093
0.059
0.003*
0.451
0.108
0.388
0.609
MS
F
P
13.07
1.44
3.09
4.96
0.50
0.64
0.28
2.68
1.22
3.95
3.03
0.89
0.81
0.88
0.183
0.375
0.009*
0.158
0.511
0.605
0.595
MS
5.39
2.32
0.32
7.64
3.29
0.23
0.44
F
3.89
5.33
1.51
12.18
25.69
0.80
1.21
Psocoptera
P
0.115
0.022*
0.231
0.020*
0.001*
0.613
0.314
MS
F
P
2.28
0.83
0.09
0.66
1.03
0.63
0.48
1.30
3.07
0.19
1.14
3.35
0.85
1.86
0.367
0.083
0.989
0.405
0.068
0.578
0.067
JBB2049_proof ■ 1 September 2010 ■ 5/12
L
D
D*L
H
H*L
H*D
H*D*L
2,4
4,8
8,16
2,4
4,8
8,16
16,32
MS
b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
Results shown for fixed main effects (L ¼ Land-use, D ¼ Distance, H ¼ Height) and their interactions; the un-replicated fields precluded testing of random effects F0 , B0 , T0 and interactions with them.
Asterisk denotes P < 0.05.
5
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
Table 2 e Comparison of the effect of land-use, distance into the cultivated areas (headland, 0 m, 5 m, 25 m, 50 m and 100 m/61 m) and heights of sticky trap (0.1 m, 1 m and
2 m) on total winged invertebrate abundance and of the nine most abundant Orders (ANOVA model 1). Orders are arranged in order of abundance on traps, with most
abundant Orders on the left.
Q1
Factor d.f.
Winged invertebrate abundance
Diptera >5 mm
Diptera <5 mm
Hymenoptera >5 mm
Hymenoptera <5 mm
Hemiptera >5 mm
586
587
588
589
590
591
592
593
594
595
596
597
598
599
600
601
602
603
604
605
606
607
608
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
640
641
642
643
644
645
646
647
648
649
650
JBB2049_proof ■ 1 September 2010 ■ 6/12
6
651
652
653
654
655
656
657
658
659
660
661
662
663
664
665
666
667
668
669
670
671
672
673
674
675
676
677
678
679
680
681
682
683
684
685
686
687
688
689
690
691
692
693
694
695
696
697
698
699
700
701
702
703
704
705
706
707
708
709
710
711
712
713
714
715
b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
Table 3 e Abundance of selected Orders within the different heights of sticky trap (0.1 m, 1 m and 2 m). Mean number of
individuals given with standard errors in brackets, reflecting variation within land-uses (willow, arable and set-aside)
between sites (n [ 3).
Order
Height
Land-use
Willow SRC
All (total abundance)
Large Diptera
Small Diptera
Large Hymenoptera
Small Hymenoptera
Large Hemiptera
Small Hemiptera
Large Lepidoptera
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
0.1 m
1m
2m
1313.74 (107.95)
1373.84 (69.43)
1367.16 (95.72)
76.02 (27.71)
62.29 (15.78)
61.86 (11.96)
27.54 (1.03)
27.25 (3.24)
27.42 (4.47)
3.70 (0.78)
5.89 (1.41)
4.60 (1.00)
32.41 (5.19)
33.31 (3.06)
36.14 (5.04)
3.51 (0.69)
3.68 (0.77)
2.73 (0.11)
3.81 (0.78)
3.64 (1.32)
3.44 (0.91)
0.70 (0.15)
0.71 (0.06)
0.66 (0.29)
diversity (Fig. 1C). Interestingly within the cultivated area
(5 m), ground flora species richness, abundance and diversity were not affected by distance, suggesting that the edge
effect is limited to within the first 5 m of the crop (species
Arable
1761.77
1299.43
985.81
22.07
15.14
20.84
57.36
40.13
29.01
1.16
0.95
0.86
20.30
15.58
12.13
1.00
0.37
0.65
2.66
1.55
1.49
0.63
0.39
0.16
(171.16)
(163.14)
(66.76)
(10.42)
(3.91)
(7.47)
(10.51)
(5.89)
(2.88)
(0.35)
(0.26)
(0.48)
(5.98)
(4.95)
(2.03)
(0.44)
(0.12)
(0.19)
(1.23)
(0.76)
(0.68)
(0.37)
(0.20)
(0.12)
Set-aside
1845.33
1205.35
900.21
58.75
37.18
21.80
65.34
42.63
28.31
2.50
1.66
0.80
14.74
11.49
9.18
3.22
1.67
1.18
4.68
2.52
2.13
2.40
0.73
0.30
(309.89)
(138.65)
(62.25)
(12.21)
(9.73)
(5.72)
(19.25)
(8.88)
(3.99)
(0.46)
(0.27)
(0.28)
(0.48)
(0.08)
(0.59)
(1.13)
(0.15)
(0.41)
(1.46)
(1.39)
(0.82)
(0.60)
(0.25)
(0.09)
richness D effect: F3,6 ¼ 1.48, P ¼ 0.311; L*D interaction:
F6,12 ¼ 0.17, P ¼ 0.986; biomass D effect: F3,6 ¼ 0.42, P ¼ 0.748;
L*D interaction: F6,12 ¼ 0.20, P ¼ 0.971; Diversity D effect:
F3,6 ¼ 1.79, P ¼ 0.249; L*D interaction: F6,12 ¼ 0.79, P ¼ 0.595).
Table 4 e Invertebrate abundance of selected Orders with distance into cultivated areas (headland, 0 m, 5 m, 25 m, 50 m and
100 m/61 m), mean number of individuals per sticky trap. Standard error is given in brackets reflecting variation within
land-uses (willow SRC, arable and set-aside) between sites (n [ 3).
Order
Distance
Land-use
Willow SRC
All (total abundance)
Small Diptera
Large Coleoptera
Thysanoptera
Headland
5m
25 m
50 m
100/61 m
Headland
5m
25 m
50 m
100/61 m
Headland
5m
25 m
50 m
100/61 m
Headland
5m
25 m
50 m
100/61 m
1934.54
1264.44
1278.26
1278.48
1002.19
54.08
23.07
22.78
20.19
16.89
2.71
0.44
0.52
0.78
0.56
3.54
0.26
0.44
0.85
0.59
(194.19)
(177.43)
(213.06)
(103.90)
(86.52)
(4.15)
(6.25)
(5.89)
(1.91)
(1.26)
(0.68)
(0.11)
(0.26)
(0.23)
(0.11)
(2.25)
(0.13)
(0.23)
(0.32)
(0.30)
Arable
1412.46 (97.35)
1187.48 (138.70)
1360.70 (104.97)
1338.83 (72.77)
1464.89 (278.23)
39.60 (4.93)
36.11 (3.26)
46.41 (7.80)
43.87 (7.21)
45.25 (11.11)
2.67 (0.95)
2.63 (0.70)
2.33 (0.72)
2.03 (0.54)
2.26 (0.70)
1.46 (1.26)
2.56 (1.84)
2.00 (1.40)
2.84 (2.31)
1.77 (0.98)
Set-aside
1280.22
1469.59
1334.05
1283.00
1217.94
42.89
51.93
47.01
41.52
43.78
4.85
3.22
3.04
2.85
3.12
1.85
2.67
2.05
2.15
2.94
(250.30)
(215.48)
(129.03)
(165.71)
(176.73)
(13.37)
(12.35)
(8.27)
(9.84)
(11.57)
(2.91)
(0.78)
(0.58)
(1.02)
(1.11)
(0.98)
(1.67)
(1.51)
(0.87)
(1.14)
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
716
717
718
719
720
721
722
723
724
725
726
727
728
729
730
731
732
733
734
735
736
737
738
739
740
741
742
743
744
745
746
747
748
749
750
751
752
753
754
755
756
757
758
759
760
761
762
763
764
765
766
767
768
769
770
771
772
773
774
775
776
777
778
779
780
JBB2049_proof ■ 1 September 2010 ■ 7/12
7
b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
Table 5 e Comparison of the effect of land-uses (willow SRC, arable and set-aside) and distance into cultivated areas
(headland, 0 m, 5 m, 25 m, 50 m and 100 m/61 m) on species richness, ground flora biomass and diversity (ANOVA model 2).
Factor
L
D
D*L
d.f.
Species richness
2, 4
5, 10
10, 20
Biomass
Diversity
MS
F
P
MS
F
P
MS
F
P
26.97
3.42
2.07
13.64
2.03
5.26
0.016
0.159
0.001
445.48
84.96
33.38
24.65
49.10
12.97
0.006
0.001
0.001
3.61
0.54
0.30
3.49
1.57
3.73
0.133
0.254
0.006
Results shown for fixed main effects (L ¼ Land-use, D ¼ Distance) and their interaction; the un-replicated fields precluded testing of random
effects F0 , B0 , T0 and interactions with them.
3.4.
Flora composition
Comparison of the most abundant plant species in willow
SRC, arable and set-aside showed that whilst some species
were present in all land-uses, differences exist in the species
composition of the three land-uses (Table 6). For example
Urtica dioica and Glechoma hederacea were found in high
abundance in willow SRC but do not feature in the top ten
most abundant species for the other land-use types (Table 6).
2
As indicated by the biomass data the mean amount of bare
ground also varied greatly with lowest levels in arable and
highest in willow SRC (Table 6).
3.5.
B
12
Plant strategies
The fraction of annual verses perennial cover was not
detectably affected by land-use (F2,4 ¼ 50.67 P ¼ 0.07).
However, a large amount of variation in the fraction of
140
120
-2
10
Ground flora biomass g 0.25m
A
N um b er o f sp e c i es p er 4m quad r at
8
6
4
2
100
80
60
40
20
0
0
Headland 0 m
C
5m
25 m
50 m 100/61 m
Headland 0 m
5m
25 m
50 m 100/61 m
1.6
1.4
Shannon diversity index
781
782
783
784
785
786
787
788
789
790
791
792
793
794
795
796
797
798
799
800
801
802
803
804
805
806
807
808
809
810
811
812
813
814
815
816
817
818
819
820
821
822
823
824
825
826
827
828
829
830
831
832
833
834
835
836
837
838
839
840
841
842
843
844
845
1.2
1.0
0.8
0.6
0.4
0.2
0.0
Headland 0 m
5m
25 m
50 m 100/61 m
Fig. 1 e Variation in the mean ground flora (A) species richness, (B) biomass and (C) diversity with land-uses (willow SRC,
arable and set-aside) and distance into the cultivated areas (headland, 0 m, 5 m, 25 m, 50 m, and 100 m/61 m). Circles
represent willow SRC, squares arable, and triangles set-aside. Error bars give standard errors, reflecting variation within
land-use between sites (n [ 3). Scale bars are not consistent.
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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847
848
849
850
851
852
853
854
855
856
857
858
859
860
861
862
863
864
865
866
867
868
869
870
871
872
873
874
875
876
877
878
879
880
881
882
883
884
885
886
887
888
889
890
891
892
893
894
895
896
897
898
899
900
901
902
903
904
905
906
907
908
909
910
JBB2049_proof ■ 1 September 2010 ■ 8/12
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911
912
913
914
915
916
917
918
919
920
921
922
923
924
925
926
927
928
929
930
931
932
933
934
935
936
937
938
939
940
941
942
943
944
945
946
947
948
949
950
951
952
953
954
955
956
957
958
959
960
961
962
963
964
965
966
967
968
969
970
971
972
973
974
975
b i o m a s s a n d b i o e n e r g y x x x ( 2 0 1 0 ) 1 e1 2
Table 6 e The ten most abundant ground flora species within each land-use (willow SRC, arable and set-aside), based on
sum cover of all quadrats percentage of bare ground also shown.
Willow SRC
Elytrigia repens
Urtica dioica
Holcus lanatus
Dactylis glomerata
A. stolonifera
Glechoma hederacea
Festuca rubra
Ranunculus repens
Agrostis capillaris
Calystegia sepium
Bare ground
% cover
Arable
% cover
Set-aside
% cover
21.5
18.3
18.3
7.9
5.3
3.9
3.8
1.9
1.8
1.6
7.7
Elytrigia repens
Bromus sterilis
Arrhenatherum elatius
Festuca rubra
Galium aparine
Fallopia convolvulus
Holcus lanatus
Polygonum aviculare
Dactylis glomerata
Lolium multiflorum
Bare ground
2.8
2.8
1.8
1.5
1.4
1.2
1.0
1.0
0.7
0.6
80.9
Holcus lanatus
Agrostis stolonifera
Taraxacum agg
Bromus hordeaceus
Bromus sterilis
A. elatius
Agrostis capillaries
Epilobium montanum
Chenopodium album
Rumex acetosella
Bare ground
13.73
6.69
5.80
5.18
3.64
3.63
3.37
2.83
2.38
2.04
23.02
annual and perennial cover was apparent in set-aside land
and especially the arable land (Fig. 2A). In contrast, willow
SRC was invariably dominated by perennial cover with mean
annual cover per sampling location never greater than 2%
(Fig. 2).
There was also a large amount of variation in life form
especially in willow SRC (Fig. 2B). Effect of distance was
present in all land-uses with increased grass cover in the
headlands of all land-uses in comparison to the cultivated
area (F5,30 ¼ 5.98 P ¼ 0.001) (Fig. 2B), but no overall effect of
land-use was detected (F2,4 ¼ 5.29 P ¼ 0.075). The large variation in life form in willow SRC reflects the patchy nature of the
flora cover in willow SRC, which both within and in particular,
between sites often alternated between either grass cover or
competitive forbs especial U. dioica (pers obs.). In contrast the
cover in arable land appears more consistent and although
not significant, the level of forb cover does increase with
distance into the cultivated area (Fig. 2B).
Competitive (Cþ) and ruderal (Rþ) establishment strategies groups are affected by land-use (Cþ F2,4 ¼ 9.53 P ¼ 0.030,
Rþ F2,4 ¼ 19.53 P ¼ 0.009) (Fig. 2C). Within these strategies
arable and set-aside land had similar levels of cover
(C F1,2 ¼ 3.72 P ¼ 0.84, R F1,2 ¼ 1.30 P ¼ 0.37) whilst willow SRC
had a higher fraction of competitive cover and an almost
complete absence of ruderal species. Competitive and ruderal
cover were also affected by distances (Cþ F5,30 ¼ 3.25
P ¼ 0.018, and Rþ F5,30 ¼ 2.69 P ¼ 0.040) with the headlands of
arable and set-aside land containing decreased ruderal and
increased competitive cover compared to the cultivated area
(Fig. 2C).
CSRþ species were present in all land-uses (Fig. 2C), with
a similar fraction of cover and no interaction with distance
present in willow SRC and set-aside (L * D F5,20 ¼ 0.51
P ¼ 0.764). In contrast in arable land, fraction of cover varied
greatly with distance, being almost absent at 100/61 m yet
accounting for over 60% of the mean cover at 25 m (Fig. 2B)
resulting in a significant interaction between land-use and
distance (F10,30 ¼ 2.16 P ¼ 0.048). However, overall CSRþ cover
was similar across all land-uses (F2,4 ¼ 0.50 P ¼ 0.640).
Stress tolerant (Sþ) species were only recorded in set-aside
land and at very low levels, accounting for only 2% 1.2% of
total cover (Fig. 2C), making testing and conclusions on the
distribution of this group inappropriate.
4.
Discussion
4.1.
Winged invertebrates
This study specifically examined invertebrate groups previously ignored in earlier studies of SRC biodiversity [4,14] and
demonstrated clear differences in the assemblage of various
winged invertebrate Orders in willow SRC compared with
arable and set-aside, particularly at canopy height. This
observation suggests that winged invertebrates in willow SRC
are associated more with the willow canopy than with the
ground flora; a finding consistent with Reddersen [32] who
also concluded that for flower-visiting insect the ground flora
within willow plantations is of little interest due to limited
flowering. For the Hymenoptera, which show increased
abundance at increased heights in willow, the attractiveness
of the canopy may be related to the food sources in terms of
leaf-feeding beetle larvae [33] and stem-feeding aphids [34].
Indeed individuals of Vespidae and Apidea families were
observed feeding on honeydew produced by aphids on willow
stem (R. Rowe pers Obs) a behaviour known for these families
[35,36]. Our data suggest therefore, that willow SRC could
provide an important resource for winged invertebrates and in
particular Hymenoptera and Hemiptera species, even if weed
control measures are increased in future as has been suggested by some plantation managers [37]. Nonetheless, we
also note here that the wider role of the ground flora in supporting invertebrate community diversity within SRC plantations requires further research. It must also be noted that
the arable fields were stubble at the time of this survey and
although this would have limited effect on the “weed” flora
recorded winged invertebrate diversity would have been
affected by the limited crop cover. However, arable fields were
expected to remain stubble or bare ploughed field for several
months [38] so comparison to arable fields in this condition
was deemed to be valid, although clearly temporal studies
thoughout the full crops cycle are needed.
The increased abundance of winged invertebrates in willow SRC headlands together with the changes in Order
abundance between the headlands and crop highlight the
importance of headlands for overall abundance and diversity.
This result supports previous work showing that the sheltered
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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Fig. 2 e Variation in the fraction (A) life history (annual or perennial), (B) life form (grass or forb) and (C) establishment
strategies (CD, CSRD, RD, SD), cover with distance. For clarity land-use are referred to by first letter, Willow SRC
represented by W, arable by A, and set-aside by S. Error bars (standard error) removed from establishment strategies for
clarity.
Please cite this article in press as: Rowe R, et al., Potential benefits of commercial willow Short Rotation Coppice (SRC) for farmscale plant and invertebrate communities in the agri-environment, Biomass and Bioenergy (2010), doi:10.1016/
j.biombioe.2010.08.046
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nature of willow headlands is beneficial to winged invertebrates [39]. There may also be an ecotone effect resulting in
the increase in invertebrate abundance and the changes in
Orders recorded.
4.2.
Ground flora
Our results illustrate the beneficial value of mature SRC
cultivation for plant community composition in the agrienvironment. In particular, we demonstrate significant variation in the primary life-history strategies exhibited by the
component plant community; i.e. SRC plantations contain
a consistently high fraction of perennial species and were
dominated by Competitive (Cþ) and Competitive e Stress
tolerant e Ruderal (CSRþ) groups, such as Holcus lanatus and U.
dioica. Although the dominance of such species is consistent
with previous studies [14,15,40], here we show a clear difference between plant community composition in SRC and the
main alternative land-use options.
The variation in plant life-history strategies between landuses is likely to reflect the reduced level of disturbance
experienced by SRC (harvesting every three years) in
comparison to the more frequent disturbance in arable and
set-aside land. As a result, willow SRC provides a more stable
habitat and consequently may play a role as a reservoir for
many components of farmland diversity. In this respect it may
provide a similar role to that attributed to arable headlands,
beetle banks, and semi-natural habitats [41e43]. The light
levels within willow SRC plantations are however likely to be
reduced in comparison to these more open habitats [15].
Indeed although no direct measure of light intensities were
taken in this study earlier studies have shown that during the
growing season photoactive radiation (PAR) is reduced by
between 98% and 88% within uncut willow plantation [15].
This is likely to affect the plant species which will successfully
establish within these plantations. Several of the dominant
plant species recorded in SRC have wider benefits for biodiversity. U. dioica for example, is host plant for a wide range of
invertebrate species including Aphididae [44] and Lepidoptera
such as Noctuidae, Nymphalidae and Pyralidae families [45],
while Dactylis glomerata is general considered a relatively high
quality grass species and is a food plant for Orthoptera species
[46] as well as Hesperidae and Satyridae larvae [45]. G. hederacea also provides a source of early spring pollen and nectar
for pollinating insects [47].
This study also clarifies the distance to which an edge
effect is apparent in willow SRC, with a consistent species
richness and ground flora biomass in the cultivated area from
5 m into the crop onwards. This suggests that whilst the crop
edge may be important in maintaining a wide range of species
most of the crop can be considered a relatively consistent
“interior” habitat.
4.3.
Implication for biodiversity and ecosystem service
Differences in ground flora species, strategies and invertebrate Order abundance between the land-uses indicate that
willow SRC can have positive benefits for farmland plant and
winged invertebrate diversity by increasing spatial and hence,
habitat heterogeneity in the landscape. Caution should be
excised however, if willow SRC is to be established on areas
with set-aside type management as this may lead to
a decrease in plant species richness and a change in species
composition.
Beyond the value of SRC for biodiversity in the agri-environment, the changes in ground flora and winged invertebrates could have wide ranging impacts for ecosystem process
and services. The increased ground cover in willow SRC may
also help to reduce soil erosion and improve water quality [4].
Whilst increase in plant species richness and the associated
leaf litter in diversity could be beneficial for soil organism
diversity, and may also affect decomposition rates [48]. The
increase in species richness and plant abundance in willow
SRC and set-aside land are also likely to have positive effects
on primary production [49] and therefore, could have important and positive effects on the abundance and diversity
within other trophic levels [50].
In the case of winged invertebrates, the increased abundance and diversity of the Hymenoptera highlights the
important role that SRC might play in ecosystem service
provision. The Hymenoptera comprise many nectivorous and
predatory species; the majority of the large Hymenoptera
caught belonged to the Vespidea with small species also
including many from the Chalcidoidea superfamily. Consequently this Order provides many species that fulfil the
important roles of pollinators and biological control agents,
services essential to continued arable crop production
worldwide [51,52].
The establishment of willow SRC plantations clearly has
the potential to increase farm-scale biodiversity and may have
particularly positive effects for Hymenoptera species and
some plant species. Careful location of these plantations
could also further maximize these positive effects on both
biodiversity and ecosystem services for example by locating
plantation in areas of high erosion risk or in arable-dominated
landscapes.
Acknowledgement
With thanks to the land owners Dave Barrett and Russell
Fraser and Fred Walter of Coppice Resource Ltd., for allowing
access to the field sites. Suzie Milner, Alex Wan and Sarah Jane
York provided field assistance. This work was funded by NERC
as part of the TSEC-Biosys consortium (NER/S/J/2006/13984) to
GT and an associated studentship to MH, GT and DG
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