B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7

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Evaluating hedgerow corridors for the conservation

of native forest herb diversity

Valérie Roy, Sylvie de Blois*

Department of Plant Science and the McGill School of Environment, McGill University, Macdonald Campus, 21, 111 Lakeshore Road,
Sainte Anne de Bellevue, Québec, Canada H9X 3V9

A R T I C L E I N F O A B S T R A C T

Article history:
Received 30 January 2007
Received in revised form
1 October 2007
Accepted 14 October 2007
Available online 26 November 2007
Keywords:
Agricultural landscapes
Biodiversity conservation
Biological connectivity
Linear habitats
Forest fragmentation
Plant dispersal
The maintenance of connecting habitats such as hedgerows in production landscapes
could become increasingly critical as species migration is expected to accelerate with
climate change. Species of particular conservation interest that could benefit from con-
necting habitats especially in agroecosystems are native forest herbs. It is still unclear,
however, which hedgerow habitats have the best potential of supporting diverse forest herb
communities, making it hard to target particular structures for conservation. Our objective
was to identify the local and landscape characteristics of hedgerows that could help predict
their potential at maximizing the richness, abundance, and diversity of native forest herbs
of temperate deciduous forests of North-East America. We used multiple regression, Mor-
an’I correlograms, and conditional autoregressive models to assess the effect of landscape
and historical variables (hedgerow age, amount of adjacent forest cover, connection to for-
ests and other hedgerows) and of local variables (hedgerow width, canopy cover, cover of
other species) on the response variables. The results show an increase in forest herb diver-
sity and abundance in hedgerows with time and with increasing nearby forest cover. Local
factors such as greater width and reduced cover of other species also relate to a greater
abundance and diversity of forest herbs in linear habitats. The fact that forest herb com-
munities can reassemble in hedgerow corridors with time implies that there could be
long-term benefits in maintaining and even creating linear habitats where there is a pool
of dispersing forest herbs.
 2007 Elsevier Ltd. All rights reserved.

1. Introduction icantly to the maintenance of biodiversity, even under

There is growing evidence that the achievement of biodiver-
sity conservation objectives at the landscape or regional scale
will have to take into account not only habitats within re-
serves, but also the contribution of habitats in production
landscapes (Saunders et al., 1991; Cabeza and Moilanen,
2003; Polasky et al., 2005). This implies a better assessment
of the ecological functions of intensively managed habitats,
including habitat corridors (Townsend and Levey, 2005; Dams-
chen et al., 2006), that would allow them to contribute signif-
suboptimal conditions. Whereas much research has been
conducted to identify key factors in the selection of optimal
habitats for reserves (e.g., Margules and Pressey, 2000; Lee
et al., 2001), there is much to be done to extend and test
similar emergent principles in habitats found in landscapes
under intense management pressure (Pereira et al., 2004;
Fischer et al., 2006).
In this paper, we investigate components of native plant
biodiversity in agroecosystems focusing on a system of woo-
dy hedgerows at the margin of agricultural fields. Because of

* Corresponding author: Fax: +1 514 398 7897.

E-mail address: Sylvie.deblois@mcgill.ca (S. de Blois).
0006-3207/$ - see front matter  2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2007.10.003
 B I O L O G I C A L C O N S E RVAT I O N
their spatial and structural attributes, hedgerow habitats can
extend forest conditions into production land (de Blois et al.,
2002b), potentially enhancing connectivity between frag-
mented forest habitats (Corbit et al., 1999). The maintenance
of connecting habitats in production landscapes could be-
come increasingly critical as species migration is expected
to accelerate with climate change. Species of particular con-
servation interest that could benefit from these habitats are
the native forest herbs of temperate deciduous forests of
North America and Europe, a subset of which has been com-
monly reported in linear habitats (Boutin and Jobin, 1998; de
Blois et al., 2002a; Freemark et al., 2002; Deckers et al.,
2005a; Roy and de Blois, 2006). It is still unclear, however,
which hedgerow habitats have the best potential of support-
ing diverse forest herb communities, making it hard to target
particular structures for conservation.
Our objective in this study was to identify the characteris-
tics of hedgerows that could help predict their potential at
maximizing native forest herb richness, abundance, and
diversity. We addressed this issue at two levels and by mod-
eling species richness, abundance, and diversity in relation
to selected predictor variables for each level of observation.
Landscape (or historical) variables such as adjacent forest
cover or hedgerow age can usually be obtained from remote
sensing data in most areas. They could help in the design of
regional or landscape conservation plans for the selection of
linear structures showing the best potential for conservation.
In our landscape models, the sampling unit was a hedgerow.
The local variables are those that depend largely on manage-
ment practices at the field scale. They could help in providing
on-site qualitative management guidelines that promote the
persistence of native species in production landscapes. In
our local models, the sampling unit was a hedgerow section
that can differ in width, canopy cover, etc. In addition, since
spatial autocorrelation can be a significant problem in a data-
set such as ours, we compared the performance of standard
regression models and models accounting for spatial covari-
ance (Lichstein et al., 2002). Finally, we discussed the insights
that such models can indirectly lead into the underlying pro-
cesses (i.e., patterns of dispersal, establishment, and sur-
vival) that help maintain the diversity of semi-natural
systems.
2. Methods
2.1. Study area
The study was conducted in an area of ca. 20 km · 20 km at
the boundary (45 40 0 N, 74 1 0 W) of Mirabel (48 594 ha) and
Deux-Montagnes (30 499 ha), two regional county municipal-
ities located west of Montreal, Quebec, Canada. This region
is part of the St. Lawrence Lowlands Ecoregion marked by
moist summers and cold, snowy winters (Ecoregions Work-
ing Group, 1989). Mature deciduous forests are dominated
by Acer saccharum Marsh, in association with Carya cordifor-
mis (Wang.) K. Koch., Fagus grandifolia Ehrh., and Fraxinus
pennsylvanica Marsh (Grandtner, 1966). The region has
undergone extensive forest clearing for agriculture, indus-
trial and urban development in the past two centuries, with
continued pressure from agricultural intensification (Jobin
1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7 299
et al., 2007) and, more recently, from suburban intensifica-
tion (pers. obs.). Forest cover in the two counties was
around 25% in 2001 and had remained stable since 1993 (Jo-
bin et al., 2007). Forest patches remaining within the agri-
cultural matrix measure 19 ha on average (Bélanger et al.,
1998). Agricultural land-uses in the two counties occupy be-
tween 42% (Deux-Montagnes) and 63% (Mirabel) of the land-
scape and include dairy farming (hay fields, pasture, fallow
lands) as well as more intensive cultivation (corn and spe-
cialized crops) (Jobin et al., 2007). The rest is urban or indus-
trial area.
2.2. Landscape variables
Using recent aerial photographs and ground survey, we se-
lected 117 hedgerows characterized by a woody cover (tall
shrub or tree) and located at the margin of agricultural fields.
We ignored hedgerows concealing a major drainage ditch be-
cause of the different type of understory vegetation in these
structures. There was no minimal distance set between
hedgerows although parallel hedgerows in this landscape will
usually be separated by a field at least 200 m wide. With the
exception of the adjacent land-use that was determined in
the field, we gathered information on historical and land-
scape-scale habitat characteristics (Table 1) through aerial
photographs. We used a series of aerial photographs from
1930 (the oldest available), 1949, 1954, 1964, 1970, 1977, 1983,
1988, 1994, and 1999 (scale from 1:20 000 to 1:5000) to evaluate
the minimum age of a hedgerow. Hedgerows in this land-
scape tend to separate properties or different types of fields.
The surveyed hedgerows originated either during the time
period covered by the aerial photographs through spontane-
ous regeneration at the edge of open fields (except for two
hedgerows that were recent forest remnants) or originated
prior to the time covered by the aerial photographs. In the
latter case, it is possible that some hedgerows present in
the 1930s were linear remnants of the forest that had origi-
nally been cleared for agriculture, but they could also have
originated at the edge of open fields. In southern Quebec, lots
tended to be large at first but were often subdivided into
smaller ones (Brisson and Bouchard, 2003), possibly resulting
in new hedgerows. As well, tree size may not correlate well
with tree age in open field conditions (de Blois and Bouchard,
1995) and therefore the exact origin of some of the hedgerows
cannot be known only from aerial photographs. We calcu-
lated minimum age of a hedgerow using the first time a woo-
dy (i.e., shrubs, scattered trees) cover was present on aerial
photographs with previous photographs showing no hedge-
row structure. We corroborated the historical data by coring
trees in hedgerow sections. The majority of trees cored were
on average younger than the age assigned to the hedgerow
structure itself. Hedgerows that had never been clear-cut
since 1930 were assigned a minimum age of 85 years old,
the age of the oldest trees.
Aerial photographs from 1999 were integrated into a GIS
(ArcView 3.3, ESRI, 2002). We calculated the total area of forest
contained in buffer zones of 50, 100, and 200 m around each
surveyed hedgerow as an estimate of the potential available
habitat for the forest species pool for recolonization in hedge-
rows (Butaye et al., 2002). These distances were chosen be-
300 B I O L O G I C A L C O N S E RVAT I O N
Table 1 – Predictor variables for landscape and local regression models (including abbreviations used in models)
Level Variable
Landscape Age of hedgerow (AGE)
Intensity of adjacent land-use (LANDUSE)
Forest area in a 50 m/100 m/200 m buffer (F50/F100/F200)
Length of the hedgerow network in a 50 m/100 m/200 m buffer (N50/N100/N200)
# of connections to a forest patch (F_CONN)
# of connections to other hedgerows (H_CONN)
Length of hedgerow*
Local Hedgerow width (WIDTH)
Canopy cover (CANOPY)
Tree basal area (TREE)
Shrub cover (SHRUB)
Ruderal herb cover (RUDERAL)
Available ground space (GROUND)
* Length was used as a covariable to control for sampling effort.
cause many species in our species pool are expected to dis-
perse mostly at small distances (Roy and de Blois, 2006). With-
in the same buffer areas, the summed length of neighboring
hedgerows was also calculated with the assumption that a
well connected network could facilitate dispersal (Sarlöv Her-
lin and Fry, 2000; Bartuszevige et al., 2006). Since longer
hedgerows generate greater buffer areas, we brought values
to a common scale by dividing forest cover by buffer area
and by dividing the total length of neighboring hedgerows
by the length of the associated hedgerow. We also noted the
number of direct connections to forest patches and to perpen-
dicular hedgerows. The adjacent land-use was classified into
three classes of land-use intensity (adapted from Boutin and
Jobin, 1998), from the lowest intensity with forage or pasture
on both sides of the hedgerow to highest intensity with cash
crops on both sides of the hedgerow.
2.3. Floristic survey
We inventoried the forest herb composition of the 117 hedge-
rows, which added up to a total of 26.6 km of linear structure
sampled. The floristic survey took place in May and June 2005
to include early spring and summer species. For this study,
‘forest herbs’ were defined as herbaceous, vascular plants
characteristic of the understory of temperate deciduous for-
est interior and described as such by Marie-Victorin (1995), a
local authority. For the total length of each hedgerow, we
positioned contiguous quadrats (5 m long by 2 m wide) cen-
tered on the midline (or close by in the presence of a stone
wall). In each quadrat, forest herb populations were charac-
terized by occurrence and abundance. We used a semi-quan-
titative measure of abundance based on the number of rooted
stems, including seedlings (class 1 = 1; class 2 = 2–10; class
3 = 11–25; class 4 = 26–50; class 5 = more than 50). For analysis
at the hedgerow level, we calculated: (1) forest herb species
richness as the total number of native forest herb species
present within a hedgerow; (2) forest herb abundance as the
sum of the abundance (using mid-class value) of all species
in all quadrats of a hedgerow, and (3) the Shannon–Weiner
P
diversity index calculated as  pi log(pi) where pi is the rela-
tive frequency of species i in the hedgerow.
1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7
Range
6–85 years
Low, medium, high
0–11 801/0–49 216/0–207 771 m2
0–275/0–1052/0–3090 m
0–2
0–4
40–590 m
1.5–32.7 m
0–100%
0–0.038 m2/m2
0–10%, 11–25%, 26–50%, 51–75%, 76–100%
0–10%, 11–25%, 26–50%, 51–75%, 76–100%
0–10%, 11–25%, 26–50%, 51–75%, 76–100%
2.4. Local variables
Once the canopy was fully developed in June–July, we re-
turned to the same hedgerows to describe habitat structure.
Starting at the same position, we sampled every four quadrats
(5 m long by 2 m wide) for the whole length of the hedgerow
for structural characteristics. Each structural quadrat is there-
fore associated with 4 contiguous floristic quadrats, which we
defined as a hedgerow section measuring 2 m · 20 m. Given
the relative homogeneity of hedgerow structure across a
given section, we assumed that the structural variables mea-
sured in the first 5 m of a section were consistent throughout
the 20 m. Consequently, for a given 20 m section, species rich-
ness, abundance and Shannon’s diversity index were calcu-
lated by pooling the vegetation data collected in the four
corresponding floristic quadrats.
For each hedgerow section, we collected information on
six local habitat variables (Table 1). We measured width be-
tween the lateral limits of woody vegetation growth. We
quantified the degree of canopy closure in the center of the
quadrat using a convex spherical densiometer (Robert E. Lem-
mon, Forest Densiometers). Total basal area characterized
tree density and was established by measuring the diameter
at breast height of each living tree greater than 3.5 cm of
diameter found in the quadrat. We visually estimated shrub
cover within the 5 · 2 m quadrat to the closest percentage
class (class 1 = 0–10%; class 2 = 11–25%; class 3 = 26–50%; class
4 = 51–75%; class 5 = 76–100%). The total cover of all herba-
ceous species other than native forest species (i.e., ruderal
herb cover) as well as the available ground space (character-
ized as ground space free from rocks, trunks or other debris)
were also visually estimated to the closest percentage class
but inside a 1 · 1 m plot positioned in the center of the larger
5 · 2 m quadrat.
2.5. Data analysis
We investigated: (1) the effect of landscape variables alone
(landscape model) and (2) the effect of local variables alone
(local model), on three separate response variables, i.e., forest
herb species richness, abundance and diversity. To insure
 B I O L O G I C A L C O N S E RVAT I O N
consistency in the way we measured hedgerow ages, we ex-
cluded from the landscape model 27 hedgerows that had a
mature structure in 1954 but for which older aerial photos
(1930, 1949) were unavailable. We also excluded the only 2
hedgerows for which we had evidence that they were still
part of a forest patch 41 years ago since they were the only
remnant hedgerows of known origin and were clearly outliers
in our system. In the landscape model, hedgerows (n = 88)
were considered the sampling unit. For the local model, sec-
tions (n = 1174) were considered the sampling unit.
We standardized all explanatory variables. We normalized
the response variables of the landscape models using a Box–
Cox transformation. Richness and abundance data of the lo-
cal models were square-root transformed in order to reduce
skewness and kurtosis. No transformation could improve
normality of local Shannon data. Because there were a signif-
icant relationship between hedgerow length and the response
variables and since we were interested in effects beyond
those related to sampling effort in hedgerows of different
length, we chose to use in our landscape models the residuals
of a linear regression with standardized hedgerow length as
the response variable (linear regression, N = 88 hedgerows,
standardized length vs richness: R2 = 0.18, p < 0.0001; length
vs abundance: R2 = 0.30, p < 0.0001; length vs diversity:
R2 = 0.12, p = 0.0015).
For the landscape and local models, we conducted three
series of analysis: (1) an ordinary least square multiple regres-
sion (OSL); (2) calculation of spatial autocorrelation in the
residuals of the preliminary regression models; (3) conditional
autoregressive model (CAR) that includes spatial effect. For
the ordinary least square (OLS) multiple regression, a step-
up procedure, as detailed below, was preferred over a step-
wise approach because some of the explanatory variables
were strongly intercorrelated (Ambrosini et al., 2002; Betts
et al., 2006). We first evaluated the relative importance of each
explanatory variable with a likelihood ratio test for nested
models (Haining, 1990; Lichstein et al., 2002; Tognelli and Kelt,
2004). Larger likelihood ratio values indicate a greater contri-
bution to the model. We manually entered independent vari-
ables in order of decreasing likelihood ratio statistic until
additional predictors became non-significant when added to
the model. The correlation structure of variables entering
final models were investigated to ensure they were not corre-
lated. For each pair of explanatory variables with a correlation
greater than 0.6, the variable with the greater residual devi-
ance was excluded.
By violating the independence assumption of classical sta-
tistical tests, spatial autocorrelation can result in an overesti-
mation of habitat effects on species distribution (Legendre,
1993; Lichstein et al., 2002; Betts et al., 2006). This effect can
be important in linear habitats (Maheu-Giroux and de Blois,
2007) and especially with a sampling scheme such as ours be-
cause nearby hedgerows or hedgerow sections are likely to
share common characteristics. We chose to retain as much
information as possible in our large dataset, including infor-
mation about spatial structure, and potentially increase
statistical power. We accounted for spatial effect statistically
by first looking for spatial autocorrelation in the residuals of
all the preliminary regression models using Moran’I correlo-
grams (Moran, 1950). For the landscape models, we set incre-
1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7 301
ments at 200 m (average field width) up to a maximum
distance of 10 000 m. For the local models, we set increments
at 20 m (length of a section) up to a maximum distance of
600 m (length of the longest hedgerow). We calculated the sta-
tistical significance of I with permutation tests (999 permuta-
tions) conducted for each distance class separately. We first
verified the global significance of each correlogram using a
regular Bonferroni correction for multiple tests (Legendre
and Legendre, 1998). We then determined the significance of
I for each distance class using the progressive Bonferroni
correction in which a is divided by the current class number
(Legendre and Legendre, 1998). These analyses were carried
out using Matlab 7.0 (Mathworks, 2004).
When spatial autocorrelation was detected, we accounted
for it using a conditional autoregressive model (CAR) (Hain-
ing, 1990; Cressie, 1993; Lichstein et al., 2002; Tognelli and
Kelt, 2004; Maheu-Giroux and de Blois, 2007). CAR models as-
sume that the response is a function of both the explanatory
variables and the values of the response at neighboring loca-
tions. Autoregressive models are similar to ordinary least
squares models, with the addition of an autocovariate, a term
related to the residual variation in neighboring locations
(Keitt et al., 2002). The neighborhood relationship is expressed
in a symmetric matrix of spatial weights. We defined weights
to decrease with increasing distance between locations
2
(wij = 1/distanceij for landscape models; wij ¼ 1=distanceij for
local models) up to a maximum neighborhood distance. The
latter was selected as the maximum distance at which the
residuals from the preliminary model were autocorrelated.
As for the preliminary models, we tested the contribution of
each explanatory variable with the presence of the autocovar-
iate using a likelihood ratio test. We used Nagelkerke R2 to as-
sess OLS and CAR models goodness of fit (Lichstein et al.,
2002). All multiple regression analyses were conducted using
R software (CRAN, 2005).
3. Results
Fifty-one forest herb species were identified in our hedgerows
(Table 2). We found between 0 and 30 forest herb species, with
an average of 10 species per hedgerow. Eight species were
rather common (frequency >50%): Maianthemum racemosum,
Actaea rubra, Viola pubescens, Circaea lutetiana, Trillium erectum,
Polygonatum pubescens, Caulophyllum thalictroides, and Viola
sororia.
For all models except the landscape model predicting
diversity, OLS residuals were positively autocorrelated, indi-
cating that the assumption of independent errors was vio-
lated and required to be corrected by the addition of an
autocovariate term. As expected, the effect of the autocovar-
iate on species occurrence or abundance was highly signifi-
cant both at the local and landscape level (Tables 3 and 4).
Although regression coefficients changed as a result of
including an autocovariate, all variables significant in the
OLS models remained significant in the CAR models. Likeli-
hood ratio statistics indicate that age and neighboring forest
cover had a similar input in the CAR landscape model and
that width was the most important variable in the CAR local
model (Fig. 1). In general, the significant variables tended to
explain abundance better than richness or Shannon diversity.
302 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7

For the landscape model, adjacent forest area and hedgerow

Table 2 – Relative frequency (number of hedgerows in
which species i occurs/total number of hedgerows) pre- age were both positively related to species richness, abun-
sented in decreasing order and relative abundance (sum dance and diversity (Table 3). Species richness also increased
of total abundance for species i in all hedgerows/sum of with the number of connections to other hedgerows and spe-
total abundance for all species) for 51 species of forest cies abundance decreased with increasing land-use intensity,
herbs sampled although these relationships were less important than the
Species Frequency Abundance one for forest cover and age. A simple linear regression of spe-
(%) (%) cies richness or species abundance on forest cover or on
hedgerow age confirms that the relationships, although weak,
1. Maianthemum racemosum (L.) Link 90.60 18.55
were highly significant (Fig. 2). The local model indicated that
2. Actaea rubra (Ait.) Willd. 85.47 6.63
3. Viola pubescens Ait. 71.79 7.11 abundance, richness and diversity increased with hedgerow
4. Circaea lutetiana L. 70.94 9.97 width, canopy cover, and available ground space (Table 4).
5. Trillium erectum L. 61.54 4.08 Shrub cover and ruderal herb cover were negatively related
6. Polygonatum pubescens (Willd.) Pursh 58.97 3.50 to the response variables, especially abundance. Tree basal
7. Caulophyllum thalictroides (L.) Michx. 53.85 1.37 area was not retained in the model. Residuals from CAR mod-
8. Viola sororia Willd. 52.14 4.99
els showed no autocorrelation suggesting that the CAR mod-
9. Maianthemum canadense Desf. 41.88 3.13
10. Athyrium felix-femina (L.) Roth 41.03 2.40
els were appropriate.
11. Hydrophyllum virginianum L. 34.19 9.85
12. Erythronium americanum Ker-Gawl. 33.33 8.12 4. Discussion
13. Sanguinaria canadensis L. 33.33 1.33
14. Trillium grandiflorum (Michx.) Salisb. 33.33 2.75 While selecting hedgerow habitats for biodiversity conserva-
15. Onoclea sensibilis L. 27.35 1.57
tion in production landscapes, two variables among the set
16. Carex radiata (Wahlenb.) Small 22.22 2.11
we considered stand out as being good predictors of forest
17. Actaea pachypoda Ell. 20.51 0.16
18. Asarum canadense L. 19.66 1.71 herbs species abundance and richness: the presence and
19. Arisaema triphyllum (L.) Schott 17.09 0.85 abundance of a forest cover in the immediate surrounding
20. Carex gracillima Schwein. 17.09 1.62 and the time since the hedgerow was first colonized by trees
21. Dryopteris carthusiana 17.09 0.48 or tall shrubs. The latter relationship suggests that the poten-
(Vill.) H.P. Fuchs tial of newly created linear habitats to support species-rich
22. Aralia nudicaulis L. 15.38 0.59
native plant communities increases with time, with clear
23. Dryopteris marginalis (L.) Gray 11.11 0.39
24. Ranunculus abortivus L. 11.11 0.38 implications for conservation and the corridor function.
25. Dryopteris intermedia 10.26 0.32 Temperate forest herbs are often characterized by long
(Muhl. Ex Willd.) Gray generation times; they produce relatively few seeds that lack
26. Matteuccia struthiopteris (L.) Todaro 10.26 1.51 adaptations for long-distance dispersal and several can repro-
27. Uvularia grandiflorum Sm. 10.26 0.42 duce vegetatively (Bierzychudek, 1982; Whigham, 2004). The
28. Allium tricoccum Ait. 8.55 0.93
maintenance of a forest cover nearby hedgerows likely pro-
29. Impatiens capensis Meerb. 8.55 0.91
vides a pool of dispersing propagules for hedgerow coloniza-
30. Thalictrum dioicum L. 6.84 0.15
31. Carex deweyana Schwein. 5.13 0.26 tion. This could be especially important for forest herbs that
32. Streptopus lanceolatus (Ait.) Reveal 5.13 0.11 have limited colonization capacities, such as ant-dispersed
33. Tiarella cordifolia L. 5.13 0.11 species and species with no specific dispersal mode. Roy
34. Prenanthes alba L. 3.42 0.03 and de Blois (2006) showed that species with these traits gen-
35. Viola canadensis L. 3.42 0.08 erally occurred less in hedgerows than in forest interior. The
36. Carex peckii Howe 2.56 0.43
dispersal limitation of forest herbs in isolated habitats or in
37. Viola labradorica Schrank 2.56 0.05
38. Carex hirtifolia Mackenzie 1.71 0.07
recent or post-agricultural forests has also been well docu-
39. Carex intumescens Rudge 1.71 0.04 mented (Peterken and Game, 1984; Grashof-Bokdam, 1997;
40. Dicentra canadensis (Goldie) Walp. 1.71 0.68 Butaye et al., 2001; Singleton et al., 2001; Kolb and Diekmann,
41. Orthilia secunda (L.) House 1.71 0.02 2004). As well, the extent of remnant (or ancient) forest cover
42. Adiantum pedatum L. 0.85 0.00 left in a landscape has been linked to the species richness of
43. Athyrium thelypterioides 0.85 0.03 forest herb communities in post-agricultural sites (Vellend,
(Michx.) Desv.
2003). Whereas hedgerows adjacent to forest patches can be
44. Carex blanda Dewey 0.85 0.02
45. Carex leptonervia (Fern.) Fern. 0.85 0.02
colonized more readily by forest species, especially if dis-
46. Carex pedunculata Muhl. ex Willd. 0.85 0.02 persal vectors use both the forest and adjacent hedgerows,
47. Dennstaedtia punctilobula 0.85 0.03 only good dispersers will colonize isolated hedgerows.
(Michx.) T. Moore At the landscape scale, plant populations are not only
48. Dicentra cucullaria (L.) Bernh. 0.85 0.02 linked by seed or propagule dispersal, but also by pollen ex-
49. Hepatica nobilis Schreb. 0.85 0.04
change. These exchanges with larger nearby populations in
50. Osmunda claytoniana L. 0.85 0.01
forest can contribute to the fitness of small connected popu-
51. Pteridium aquilinum (L.) Kühn 0.85 0.05
lations in hedgerows, especially for populations of self-
Note: Nomenclature follows the Integrated Taxonomic Information incompatible species such as Trillium grandiflorum (Schmucki
System (ITIS) on-line database (http://www.itis.usda.gov).
and de Blois, in review). Populations of this species have been
 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7 303

Table 3 – Parameter estimates with significance level for Table 4 – Parameter estimates with significance level for
two types of regression models (OLS and CAR) relating two types of regression models (OLS and CAR) relating
each of forest herb richness, abundance, or diversity in each of forest herb richness, abundance, or diversity in
hedgerows with landscape variables hedgerow sections with local variables
Variable OLS modela CAR modelb Variable OLS modela CAR modelb

Richness Richness
AGE 0.38* 0.32* WIDTH 0.11**** 0.14****
F200 0.46** 0.54**** CANOPY 0.07* 0.05*
H_CONN – 0.24  SHRUB 0.16**** 0.09****
Autocov 200 m – 25.72*** RUDERAL 0.14**** 0.07***
GROUND 0.17**** 0.07****
Abundance
Autocov 280 m – 131.53****
AGE 1.23* 2.20****
F200 1.64** 1.44** Abundance
LANDUSE – 1.03* WIDTH 0.68** 0.93****
Autocovariate 400 m – 51.91**** CANOPY 0.37  0.26 
SHRUB 1.08**** 0.66****
Diversity
RUDERAL 1.30**** 0.69****
AGE 0.16* N/Ac
GROUND 1.54**** 0.72****
F200 0.21** N/A
Autocov 500 m – 138.40****
Only the significant variables are included. Refer to Table 1 for
Diversity
definitions of abbreviations.
WIDTH 0.04**** 0.05****
a OLS = ordinary least squares model based on landscape
CANOPY 0.02* 0.02*
variables. The model does not account for spatial covariance.
SHRUB 0.05**** 0.03****
b CAR = conditional autoregressive model including an autoco-
RUDERAL 0.04**** 0.02**
variate term that accounts for spatial effect up to the maximum
GROUND 0.05**** 0.02**
distance at which the residuals from the preliminary model were
Autocov 500 m – 128.54****
autocorrelated.
c N/A = absence of autocorrelation in OLS model based on diver- Only the significant variables are included. Refer to Table 1 for
sity makes correction with a CAR model unnecessary. definitions of abbreviations.
* p 6 0.01. a OLS = ordinary least squares model based only on local variables.
** p 6 0.001. The model does not account for spatial covariance.
*** p 6 0.0001. b CAR = conditional autoregressive model including an autoco-
**** p 6 0.00001. variate term that accounts for spatial effect up to the maximum
  p 6 0.05. distance at which the residuals from the preliminary model were
autocorrelated.
* p 6 0.01.
** p 6 0.001.
found to be more genetically diverse in large forest popula- *** p 6 0.0001.
tions than in small populations (Vellend, 2004), and could be **** p 6 0.00001.
more diverse in hedgerows connected through pollen   p 6 0.05.
exchanges to large forest populations than in isolated hedge-
rows. In our system and at the temporal and spatial scale of
our observations, the beneficial effects of connectivity seem dispersal limitation (Ehrlén and Eriksson, 2000; Verheyen
to depend less on the maintenance of one or more direct con- and Hermy, 2004). Our oldest hedgerows are of uncertain ori-
nections with forests, although connections to other hedge- gin and may have included remnants that would contribute
rows seem to play a role, than on landscape context and the to high diversity because they retained species from the origi-
extent of nearby forests. Increasing the number of intersec- nal forest pool. The only two known remnant outliers that we
tions in a hedgerow network could result in higher localized sampled were clearly species-rich, but were also young
plant diversity or abundance at the nodes because of in- hedgerow structure that may have yet to lose the species
creased width or increased activities of animals dispersing most sensitive to edge effects (‘extinction debt’ sensu Tilman
seeds (Deckers et al., 2005b). et al., 1994). Nevertheless, the relationship we measured be-
Because of the traits of forest herbs, we expect a time-lag tween hedgerow age and species richness held even when
in their response to changes in the landscape, either after we considered only a subset of the youngest hedgerows that
the fragmentation of continuous forest into remnant linear have clearly originated during the time period we considered
habitats or when new habitat becomes available through (n = 49; p < 0.001). Since the absence of a seed bank in most
the formation of spontaneous hedgerows. Some species with forest herb species (Hyatt and Casper, 2000) implies a reliance
long-life cycle may persist well after fragmentation and their on propagule dispersal to colonize new sites, this result and
presence may reflect more past than current landscape condi- the observed influence of adjacent forest cover suggest that
tions (Lindborg and Eriksson, 2004; Vellend et al., 2006). On hedgerows are indeed colonized by a portion of the forest spe-
the other hand, secondary or post-agricultural forests often cies pool over time. Colonization appears to occur at relatively
remain species-poor for extended periods of time in spite of short (i.e., <30 years) time scale for a subset of ‘fast’ species
site suitability (Honnay et al., 1999; Bellemare et al., 2002; that could benefit from the restoration or creation of connect-
Flinn and Vellend, 2005; Helm et al., 2006), likely because of ing habitats, especially when the need to migrate increases as
304 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7

Ordinary least squares

Richness (OLS)
80
Conditional

Likelihood ratio statistic

70
autoregressive (CAR)
60

50

40

30

20

10

0
Age F200 H_conn Width Canopy Shrub Ruderal Ground

Abundance
80
Likelihood ratio statistic

70

60

50

40

30

20

10

0
Age F200 Landuse Width Canopy Shrub Ruderal Ground

Diversity
80
Likelihood ratio statistic

70

60

50

40

30

20

10

0
Age F200 Width Canopy Shrub Ruderal Ground
Significant explanatory variables

Fig. 1 – Likelihood ratio (LR) statistics for predictor variables in ordinary least squares (OLS) and conditional autoregressive
(CAR) models to explain the richness, abundance, and diversity of forest herbs in hedgerows. Larger LR values indicate
greater contribution to model fit. Only significant variables were included. The dotted lines separate the variables from the
landscape model (left) from those of the local model (right). Refer to Table 1 for definitions of abbreviations.

expected with global warming. It will take much more time,
however, for a particular hedgerow to acquire a significant
proportion of the forest species pool, including the slow mi-
grants, and potentially contribute to metapopulation dynam-
ics if exchanges (either of seeds or pollen) are maintained
with adjacent forest patches. This also assumes that manage-
ment practices within the hedgerows are adequate to allow
populations to survive and migrate at significant temporal
scale. In this study, we did not consider species composition
or traits. A previous study investigating functional traits in
hedgerows of the same landscape, however, showed that spe-
cies with poor dispersal capacity and species that flower very
early in the season were less likely to be present or abundant
in hedgerows compared to forest interiors (Roy and de Blois,
2006), suggesting that both dispersal effect and niche effect,
perhaps at different temporal scale (Verheyen et al., 2003),
act on species distribution in this system.
Among the local habitat variables, hedgerow width had
the largest independent contribution and demonstrated a
positive relationship with all three response variables, but
mostly with species abundance. Hedgerows have a high
edge-to-area ratio and are consequently more exposed to
fluctuations in temperature and moisture than forests
(Murcia, 1995; Roy and de Blois, 2006). For organisms adapted
 B I O L O G I C A L C O N S E RVAT I O N 1 4 1 ( 2 0 0 8 ) 2 9 8 –3 0 7 305

8 8
R2 = 0.13
7 R2 = 0.16 7 p = 0.0004
Species richness

Species richness
6 p < 0.0001 6
5 5
4 4
3 3
2 2
1 1
0 0
-2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 -2 -1 0 1 2 3 4

35
30
2
R 2 = 0.10

Species abundance
30
Species abundance

R = 0.14 p = 0.0024
25
p = 0.0004 25
20
20
15 15
10 10
5 5

0 0
-2.5 -2 -1.5 -1 -0.5 0 0.5 1 1.5 2 -2 -1 0 1 2 3 4

Hedgerow age Forest cover 200m buffer

Fig. 2 – Scatterplots of total species richness and total species abundance (a) with hedgerow age, and (b) with forest cover in a
200 m buffer. Species richness and abundance were normalized using a Box–Cox transformation. Hedgerow age and forest
cover were standardized. N = 88 hedgerows used in the landscape models.

to a more uniform environment such as forest herbs, habitat
suitability is likely improved in wider hedgerows by the pres-
ence of a core area characterized by more buffered, temperate
habitat conditions, at least at the scale at which forest herbs
perceive their environment. In fact, more forest herbs oc-
curred in the center of hedgerows than at the edge (unpub-
lished). In addition, given that some forest herbs in our
species pool are animal-dispersed, hedgerow width could
indirectly affect plant populations by influencing animal use
of the linear network. In Sweden, Sarlöv Herlin and Fry
(2000) demonstrated that the number of animal-dispersed
woody species increased more significantly with hedgerow
width than the number of wind-dispersed species. By provid-
ing more resources and less exposure to predators, wide
hedgerows generally favour the presence of birds (Hinsley
and Bellamy, 2000) and possibly of small mammals.
Within hedgerows, the abundance of forest herbs may be
limited by competition from shrubs and ruderal species that
benefit from increased disturbances related to more intense
field management. In another hedgerow system of a similar
landscape, it was found that, independent of hedgerow age,
competitive ruderal species were more likely to colonize nar-
row hedgerows adjacent to the more intensively managed
fields, whereas the probability of finding forest herbs in
hedgerows increased as the intensity of adjacent agricultural
practices (from intensive crop to fallow land) decreased (de
Blois et al., 2002a). The microclimatic conditions of hedge-
rows generally characterized by increased light penetration,
increased air and soil temperature fluctuations, decreased
air humidity and an increased level of agronutrients in the
soil give a competitive advantage to ruderal herbaceous spe-
cies and invasive shrub species over shade-tolerant forest
interior species (Honnay et al., 2005; Roy and de Blois, 2006).
Forest herbs in some hedgerows are subjected to levels of
shrub competition not experienced in forest patches. A dense
shrub cover can change the seasonal patterns of light avail-
ability and affect understory herbs that depend on the period
before canopy expansion for much of their annual carbon
gain (Miller and Gorchov, 2004). The resulting reduction in
growth and fecundity can suppress the long-term reproduc-
tive success of herbaceous species and consequently reduce
population growth rate.
5. Conclusion
The potential for hedgerows to support relatively diverse na-
tive forest herb communities increases with time and the
maintenance of a forest cover in the adjacent landscape,
and to a lesser extent, with connections to other hedgerows.
Management practices that lead to wider hedgerows with re-
duced competition from other species, by minimizing, for in-
stance, disturbances from field maintenance, are expected to
result in an increase in the richness and abundance of forest
herbs within linear habitats. A direct link to a forest patch
does not appear to influence forest herbs in this system, but
it may possibly affect which species will be able to colonize.
The fact that forest herb communities, or at least a subset
of the regional pool, can reassemble in hedgerows with time
implies that there could be long-term benefits in maintaining
and creating linear habitats where there is a pool of dispers-
ing forest herb propagules. A structurally diverse hedgerow
flora can also have a positive effect on the diversity of other
306 B I O L O G I C A L C O N S E RVAT I O N
taxa (Jobin et al., 2001). Most hedgerows, however, are subject
to disturbances from agricultural management practices and
even species that have managed to colonize and establish
small populations can be extirpated. With inappropriate
management, hedgerows may act more as sinks than as valu-
able biodiversity conservation units, but the conflict between
the legitimate need to maintain weed and pest-free agricul-
tural fields and the urgency to conserve native species where
they are most threatened is not easily resolved. We believe
that hedgerows can contribute, at least for a subset of species,
to extend forest conditions into the agricultural matrix, possi-
bly mitigating in part the impacts of habitat loss and frag-
mentation on dispersing species with minimal effect on
crop yield. Based on our results, we should at the very least
aim to conserve the remaining species-rich hedgerows and
facilitate, whenever conditions allow, the emergence of new
linear structures that will have the potential to sustain the
dispersal of native plants in the long-term.
Acknowledgments
This research was supported by an FQRNT scholarship to V.R.
and NSERC and FQRNT grants to S.d.B. We are grateful to J.
Snider, M. Bourgon-Desroches and V. Paul for their assistance
in the field and to R. Schmucki for help with statistical anal-
ysis. We wish to thank E. Bennett, M. Vellend, M. Waterway
and three anonymous reviewers for commenting on an ear-
lier draft of this manuscript.
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