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22001133 NORTNHorEthAeSaTsEteRrNn NNaAtTuUraRliAstLIST 2V0(o4l). :2609,4 N–7o0. 94
Plant Communities within Atlantic Coastal Heathlands in
Nova Scotia
Robert P. Cameron1,* and Soren Bondrup-Nielsen2
Abstract - Coastal heathlands are rare ecosystems that provide habitat for rare species in
Nova Scotia. Thirty-nine plots were established in Nova Scotia heathlands to assess plant
community composition and occurrence of rare plants. Analysis of species richness and
multidimensional scaling (MDS) revealed that heathland communities are varied, with
differences between regions, inland and coastal sites, and between physiognomy types.
Six rare plants occurred within 9 of 39 plots. Coastal heathland communities were found
to have greater species richness and variation in community type than previously thought.
Heathland rare plants are not restricted to any particular community type; rather, rare
coastal plants in Nova Scotia occur in a wide variety of community types. Coastal heathlands
add diversity to the mostly forested landscape of Nova Scotia and provide habitat
for rare species.
Introduction
Coastal heathlands occur as treeless or nearly treeless dwarf-shrub communities
on headlands or exposed land along the Atlantic coast of southeastern Canada and
northeastern US (Dunwiddie et al. 1996). Exposure to salt spray and winds from the
North Atlantic are key factors in maintaining most coastal heathland communities
(Griffiths 2006; Griffiths and Orians 2003, 2004). Some heathland communities may
also benefit from human disturbance such as burning or livestock-grazing, which
help maintain their community composition (Dunwiddie 1990).
Coastal heathlands provide habitat for rare plants in the northeastern US (Clarke
and Patterson 2007, Dunwiddie 1990) and Nova Scotia (Pronych and Wilson 1993).
Oberndorfer and Lundholm (2009) found rare species richness was greater where
heathland vegetation height is low. Rare plants reported from coastal heathlands
in the northeastern US also occur in low-shrub communities (Dunwiddie 1990,
Godfrey and Alpert 1985, Noss et al. 1995). Most of the rare plants known from
heathlands have an arctic-alpine distribution and are at the southern extent of their
ranges in southeastern Canada and northeastern US. Arctic-alpine plants may be
able to persist in coastal heathlands because harsh conditions limit the establishment
of more competitive species.
Coastal heathlands make up a small area of Nova Scotia and are considered a
rare community type (Cameron et al. 2010b), similar to the situation in the northeastern
US (Noss et al. 1995). Conservation efforts have been in place in the US for
at least 25 years (Godfrey and Alpert 1985), but only recently has the importance of
coastal heathlands in Nova Scotia been recognized. For example, the passing of an
1Nova Scotia Environment, Protected Areas Branch, PO Box 442, Halifax, NS B3J 2P8
Canada. 2Centre for Wildlife and Conservation Biology, Acadia University, Wolfville, NS
B4P 2R6 Canada. *Corresponding author - camerorp@gov.ns.ca.
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amendment to the Off Highway Vehicles Act (1989) in 2005 specifically prohibits
OHV use in coastal heathlands.
The Nova Scotia Department of the Environment is charged with designating
and managing provincial protected areas. These areas are selected to protect
rare and endangered species and communities. Given the rarity of heathland
communities and their significance in containing rare species, it is imperative to
document the community composition and occurrence of rare plants within coastal
heathland communities.
Methods
Sampling
Provincial wilderness areas and nature reserves make an ideal network for the
study of community structure and function because these lands represent the variety
of ecosystems present in the province, and they are relatively undisturbed by human
impacts. Wilderness areas are legally protected from development activities including
forestry, mining, and road building (Cameron 2004).
The Nova Scotia Department of the Environment is involved in a larger project
to assess the biodiversity of these protected wilderness areas and nature reserves in
Nova Scotia; the data used in this study were selected from a more extensive dataset
based on a plot design. Methods for plot selection were designed following the Ecological
Society of America Guidelines for Describing Associations and Alliances of
the US National Vegetation Classification (Jennings et al. 2004).
Data were collected from 29 wilderness areas and 6 nature reserves. We identified
topographic features and dominant plant communities within each of these
areas using Nova Scotia Environment Geographical Information System Ecosystem
Classification (Cameron and Williams 2011). A transect was placed within each area
such that it traversed the variety of topographical features and dominant plant communities
of each landscape. We walked these transects and established plots along
or near these lines in areas that represented a relatively homogenous vegetation
community. We established new plots each time we encountered a different homogenous
vegetation community. Using criteria from Jennings et al. (2004), we defined
a homogeneous vegetation community as “contiguous areas of vegetation that are
reasonably uniform in physiognomy, floristic composition, and environment”. Following
protocols of Jennings et al. (2004), we conducted a reconnaissance of each
vegetation community encountered to determine the extent and degree of variation
of plant species, plant physiognomy, and environmental gradients within the
community. Environmental gradients can include a variety of factors, but we only
collected data on soil drainage and soil depth. We subjectively selected a location
within each vegetation community that best represented the variation in species,
physiognomy, and environmental gradients, and established a 20-m x 20-m plot.
Based on physiognomy and species composition, we identified heathlands from
the larger dataset of plots. Plots were considered as heathland if they had less than
25% tree cover in the canopy layer, were dominated by ericaceous plants or lichens,
and had well- to imperfectly-drained, but not saturated soils that were dominated
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2013 Northeastern Naturalist Vol. 20, No. 4
by facultative- or obligate-wetland plant species (Davis and Browne 1994). From
the larger dataset of 29 Wilderness Areas and 6 Nature Reserves, we selected 39
plots of heathlands from 8 Wilderness Areas and 2 Nature Reserves to be included
in this study (Fig. 1). Protected areas selected for this study were Blue Mountain-
Birch Cove Lakes, Bowers Meadows, Canso Coastal Barrens, Gabarus, Scatarie
Island, Ship Harbour-Long Lake, Tidney River, and Tobeatic Wilderness Areas, and
Blandford and Duncan’s Cove Nature Reserves. We sampled an additional 3 plots
in Baleine and Kelly Long Lake, undesignated crownlands, because rare plants had
been documented from these heathlands.
We collected presence and abundance of plant species, and data regarding environmental
site factors at each plot following standards set out by the Ecological
Society of America (Table 1; Jennings et al. 2004). We recorded cover-abundance
classes for each plant species by layer within each plot using seven cover-classes
that were based on the percentage of ground covered by each species within the 20-m
Figure 1. Heathland plant-community study-site locations (white dots) in Nova Scotia,
Canada. Numbers refer to the following study sites: 1 = Blue Mountain-Birch Cove Lakes
(1 plot), 2 = Bowers Meadows (2 plots), 3 = Canso Coastal Barrens (4 plots), 4 = Gabarus
(4 plots), 5 = Scatarie Island (6 plots), 6 = Ship Harbour-Long Lake (1 plot), 7 = Tidney
River (4 plots), 8 = Tobeatic (9 plots) wilderness areas, 9 = Duncan’s Cove Nature Reserve
(1 plot), 10 = Blandford Nature Reserve (3 plots), 11 = Baleine (2 plots), and 12 = Kelly
Long Lake (2 plots).
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x 20-m plot: 0 = absence, 1 = trace (less than 0.1%), 2 = 0.1–1%, 3 = 1–5%, 4 = 6–25%, 5 =
26–50%, 6 = 51–75%, 7 = 76–100%. The four layers in which we estimated plant
cover classes were: canopy, shrub, herb, and moss/lichen. We assessed cover classes
for each layer independently of other layers; thus, species cover can exceed 100%
if more than one layer is summed. We used guidelines provided by Jennings et al.
(2004) suggesting that layers be defined in the field using growth form. For example,
we considered an individual plant in a plot to be in the shrub layer if that plant was
within the range of heights commonly observed for the region’s mature shrub species.
Within a plot, we listed each species occurring in every layer in which it was
observed, with a separate cover estimate for its abundance in each of these layers. For
example, a tree species could potentially be recorded in herb, shrub, and canopy if
seedlings, saplings, and mature trees were present in the plot.
We designated plots as high shrub when heath species dominated the shrub
layer, or low shrub when heath species were largely absent from the shrub layer
but occurred in the herbaceous layer. We considered heath species to be those species
known to dominate heathland communities as suggested by Davis and Browne
(1996). We also divided our samples geographically: we designated plots south and
west of the City of Halifax as western, and plots north and east of the City of Halifax
as eastern. We considered coastal plots as those occurring within the Atlantic
Coastal Climate Region of Nova Scotia as designated by Dzikowski (1985). Coastal
plots included all areas except Tobeatic Wilderness Area, which was the only inland
site and contained all the inland plots used in our analysis.
Analysis
We used Kruskal-Wallis nonparametric analysis to compare species richness
among heathland types, and multidimensional scaling (MDS) to assess differences
among categories of heathlands. MDS is one of several multivariate ordination
methods that can be used to arrange communities along environmental gradients
based on community composition (ter Braak 1987); differences (or similarities)
between communities are calculated and then plotted so that the distances between
sites are maximally correlated with ecological distances.
Results
Thirty plots occurred in coastal heathlands, and 9 plots occurred in inland
heathlands. Of the 9 inland plots, 2 were low shrub and 7 were high shrub. Of the
Table 1. Environmental site factors collected in 39 heathland plots in Nova Scotia, Canada.
Environmental site factor Assessment criteria
Aspect Direction plot faces, measured in degrees on azimuth compass
Slope Degree of steepness, measured as percent slope
Elevation Measured in meters above sea level
Soil drainage How well water is carried away from the site, categorized as: excessive,
well, imperfect, poor, or saturated
Topographic position Categorized as: crest, upper slope, middle slope, lower slope, toe slope,
flat, depression, or floodplain
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coastal heathlands, 20 were low shrub and 10 were high shrub. The majority (18)
of coastal heathland plots were in the eastern region of the province; the remaining
9 plots were in the western region. We recorded 102 species of vascular plants and
37 bryophyte and lichen species in our plots (Appendix 1).
Community types
MDS showed good separation of community types using all species (Fig. 2a),
shrubs by themselves (Fig. 2b), and shrubs and herbs together (Fig. 2c), but not
when we tested herbs or non-vascular plants.
High-shrub plots were located on the left side of the MDS plot, and low-shrub
plots were found on the right side of the MDS plot. Differences in plant species
presence and cover classes are clearly evident between high- and low-shrub communities.
Although many of the high-shrub species were present in low-shrub
communities, they occurred at much lower cover classes there than in the highshrub
communities. However, many species occurred in the low-shrub communities
that did not occur in the high-shrub type; some examples include Corema conradii
(Broom Crowberry), Empetrum nigrum (Black Crowberry), Juniperus communis
(Common Juniper), Sibbaldiopsis tridentata (Three-toothed Cinquefoil), Cladonia
maxima (Asahina) Ahti (Giant Cladonia), C. stellaris (Star Reindeer Lichen) and
C. boryi (Bory’s Cup Lichen).
Differences in community composition between eastern and western plots were
more subtle; however, eastern plots generally occurred in the upper right of the
MDS plot. Many of the plant species occurred in both eastern and western plots,
Figure 2. Multidimensional scaling graphs of 39 heathland plant-community plots in Nova
Scotia. Plots are shown as symbols: crosses are inland high-shrub, downward-facing triangles
are western coastal high-shrub, squares are eastern coastal high-shrub, upward-facing
triangles are eastern coastal low-shrub, stars are western coastal low-shrub, and circles are
inland low-shrub. A = all species, B = shrubs, and C = shrubs and herbs.
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but they were observed in different cover classes and they differed in frequency of
occurrence in plots. For example western plots had a greater frequency of Pteridium
aquilinum (Bracken Fern), Rhododendron canadense (Rhodora) and Trientalis borealis
(Star Flower). Eastern plots had a greater frequency of Photinia melanocarpa
(Black Chokeberry), Myrica gale (Sweet Gale) and Cladonia rangiferina (Greygreen
Reindeer Lichen).
Two coastal plots (plots 16 and 17) and one inland plot (plot15), had species
compositions different from all other plots. The two coastal plots occurred in the
Canso Coastal Barrens Wilderness Area in Guysborough County in eastern Nova
Scotia. Plot 16 was dominated by Deschampsia flexuosa (Common Hair Grass) and
Carex trisperma (Three-seeded Sedge). Plot 17 was dominated by Osmunda cinnamomea
(Cinnamon Fern) with a mix of common herbs such as Aralia nudicaulis
(Wild Sarsaparilla), aster sp., Clintonia borealis (Blue Bead Lily) and Prenanthes
trifoliolata (Dwarf Rattlesnakeroot). Inland plot 15 had very low species richness
with large areas of exposed bedrock.
Low-shrub coastal communities. Low-shrub coastal heathlands were most often
dominated by Black Crowberry. Where Black Crowberry was not dominant, low
coastal heathlands tended to be dominated by Broom Crowberry. Ledum groenlandicum
(Labrador Tea), Maianthemum canadense (Canada Mayflower), and
Three-toothed Cinquefoil were often found as well, but with lower cover than either
species of Crowberry. Star Reindeer Lichen and Bory’s Cup Lichen were also frequently
found along with other Cladonia spp. (reindeer lichens). Low-shrub coastal
communities were found either on headlands, exposed ground, or in protected locations
on rocky ground with little or no soil.
High-shrub coastal communities. High-shrub coastal heathlands were dominated
by a wide mix of ericaceous species. Kalmia angustifolia (Lambkill) was most
often found, but Labrador Tea, Vaccinium angustifolium (Lowbush Blueberry), and
Rhodora were also frequently found. The high-shrub heathland usually occurred at
least several hundred meters inland from headlands and exposed areas.
Tobeatic inland shrub communities. One plot had little cover in the shrub layer
and was dominated by Broom Crowberry and Arctostaphylos uva-ursi (Bearberry).
Grey-green Reindeer Lichen was common in the moss/lichen layer in this plot. All
other plots in the Tobeatic heathland were dominated by ericaceous shrubs. Gaylussacia
baccata (Black Huckleberry) and Ilex glabra (Inkberry) were abundant, but
Lambkill and Rhodora were also frequently found in plots. The herb layer was often
dominated by Bracken Fern.
Species richness
Coastal high-shrub communities in the east had the greatest species richness,
likely because of the high number of species in the shrub layer (Table 2). Three
species found only in the coastal high-shrub communities in the east include Abies
balsamea (Balsam Fir), Betula papyrifera (White Birch), and Sorbus americana
(Mountain-ash). Coastal low-shrub communities in the west however, also had
high species richness. The relatively high species richness of coastal low-shrub
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2013 Northeastern Naturalist Vol. 20, No. 4
communities in the west was a reflection of the high species diversity in the herb
and non-vascular layers. There were 13 species in the herb layer and 4 species
in moss/lichen layer that were found only in the coastal low-shrub communities in
the west. Inland high-shrub and inland low-shrub had the lowest and second lowest
species richness, respectively.
Rare species
Six species of rare plants occurred in 9 plots (Table 3). Vaccinium uglinosum
(Alpine Whortleberry) occurred in well-drained hummock-tops or mounds. All Alpine
Whortleberry sites had high Black Crowberry cover and occurred in low- and
high-shrub communities. The high shrub plots had high cover of Black Huckleberry
and Morella pensylvanica (Bayberry).
Minuartia glabra (Mountain Sandwort) occurred on exposed bedrock with high
lichen and moss cover. Star Reindeer Lichen, Bory’s Cup Lichen, and Andreaea
rupestris (Andreaea Moss) were common in Mountain Sandwort plots. Broom
Crowberry and Pinus banksiana (Jack Pine) were also frequent at these sites.
Betula michauxii (Michaux’s Dwarf Birch) occurred in imperfectly to poorly
drained sites, sometimes on the edge of ponds in both high- and low-shrub communities.
Sphagnum spp. (peat mosses) cover, frequently including Sphagnum
magellancium (Magellan’s Sphagnum), was high at Michaux’s Dwarf Birch sites.
Common shrub species were Lambkill and Bayberry.
Vaccinium boreale (Northern Blueberry) occurred with Black Crowberry and
Alpine Whortleberry on well-drained, exposed headlands. Alnus viridus (Downy
Alder) was common, and Common Juniper and Lambkill were also found in
these plots.
Dwarf Rattlesnakeroot occurred on well-drained sites, mostly low-shrub headlands,
often with Black Crowberry. Common juniper, Broom Crowberry, and
Table 2. Comparisons of mean and standard deviation of species richness, including sample size (n)
within four layers of vegetation (canopy, shrub, herb, and non-vascular layers, as well as total), using
the Kruskal-Wallis non-parametric test showing chi-square values and probability (P) for 6 shrub
habitats in Nova Scotia; low = low-shrub, and high = high shrub.
Coastal Inland
Low High Low High Low High Chi
east east west west west west squared P
n 14 4 6 6 2 7
Canopy 0.36 0.00 1.50 1.33 1.00 1.00 10.48 0.06
1.08 0.00 1.97 0.82 1.41 1.15
Shrub 3.86 10.50 1.83 6.33 5.00 6.00 12.61 0.03
2.82 5.08 2.79 2.58 2.82 2.58
Herb 8.21 7.50 9.33 6.33 3.5 1.71 17.06 0.00
3.49 4.65 5.43 2.88 0.71 0.95
Non-vascular 3.21 5.50 7.50 2.17 2.50 1.71 13.62 0.02
1.89 3.32 3.93 2.86 2.12 1.25
Total 15.64 23.50 20.17 16.17 12.00 10.43 11.51 0.04
4.22 11.09 8.77 5.11 5.66 3.26
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Vaccinium vitis-idaea (Foxberry) were often found in the same plots. Rare species
found in heathlands outside plots included Cornus suecica L. (Lapland Cornel) and
Schizaea pusilla Pursh (Curlygrass Fern) on Scatarie Island, and Solidago multiradiata
Aiton (Rocky Mountain Goldenrod) in Baleine.
Discussion
Community types
Few empirical studies have been made of Nova Scotia coastal heathlands. This
study and work by Oberndorfer and Lundholm (2009) suggest that coastal heathland
communities may be more complex than first thought. Coastal heathland
communities are clearly different from the inland heath communities found in
the Tobeatic Wilderness Area. Analysis using MDS showed a definite separation
between inland and coastal communities. Davis and Browne (1996) also suggest
a difference between inland and coastal heathlands. This study found Black
Crowberry and Empetrum eamesii Fernald &Wiegand (Red Crowberry) to be key
indicators of coastal heathlands, a finding also suggested by Davis and Browne
(1996). Although Davis and Browne (1996) suggest Huckleberry, Kalmia polifolia
(Bog Laurel), Bearberry, and Rhodora are indicators of inland heath, we frequently
recorded them in our coastal heathland plots. Oberndorfer and Lundholm (2009)
also found these species in many of their coastal barren plots.
Table 3. Rarity ranking, locations where found, and number of plots found for rare plant species in 9
of 39 plots in Nova Scotia heathlands. NS ranking = Nova Scotia provincial ranking
Number
NS ACCDC plots
Species rankingA rankingB Locations found found
Alpine Whortleberry Yellow S2 Scatarie Island 2
(Vaccinium uliginosum L.)
Dwarf Rattlesnakeroot Yellow S5C Canso Coastal Barrens, 5
(Prenanthes trifoliolata (Cass.) Fernald) Gabraus, Scatarie Island
Michaux’s Dwarf Birch Yellow S2 Baliene, Kelly Long Lake 2
(Betula michuaxii Spach)
Mountain Sandwort Yellow S2D Blandford, Blue Mountain 4
(Minuartia groenlandica (Retz.) Ostenf.) Birch Cove Lakes
Northern Blueberry Red S2 Scatarie Island 2
(Vaccinium boreale I.V. Hall & Aalders)
Red Crowberry Yellow S2/S3 Duncan’s Cove 1
(Empetrum eamesii Fernald & Wiegand)
ARed = known or thought to be at risk; Yellow = sensitive to human activities or natural events.
BAtlantic Canada Conservation Data Centre (ACCDC): S2 - Rare (May be vulnerable to extirpation
due to rarity or other factors, 6 to 20 occurrences or few remaining individuals); S3 - Uncommon, or
found only in a restricted range, even if abundant at some locations (21 to 100 occurrences).
CHad been known as Prenanthes nana, but P. nana is no longer recognized as a distinct species from
P. trifoliolata by ACCDC.
DRecent discoveries of large populations of this species in southern Nova Scotia will likely result in
this species receiving a lower rank (S. Blaney, Atlantic Canada Conservation Data Centre, Sackville,
NB, Canada, pers. comm.).
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Lichens may be another indicator of coastal heathlands. We found Bory’s Cup
Lichen in 14 of 30 coastal plots and Cladonia terrae-novae in 12 of 30 coastal
plots; we found neither species in inland plots. Oberndorfer and Lundholm (2009)
found the lichen community in their coastal plots was the most consistent of any
group among plots. Cameron et al. (2010a) also suggested these species as possible
coastal community indicators.
MDS analysis suggests a distinct difference between high-and low-shrub coastal
heathlands (Fig. 2a). Strang (1971) also found distinct high- and low-shrub communities
in inland heathlands in Nova Scotia. Unlike inland high-shrub communities,
however, species with a coastal affinity such as Downy Alder and Bayberry have
high cover in coastal high-shrub. Strang (1971) described low-shrub communities
as occurring on dry hummock-tops and high-shrubs on slopes and depressions. In
our study of coastal heathlands, low-shrub communities dominated headlands and
exposed nearshore areas. High-shrub communities tended to occur in less exposed
areas, often several hundred meters from the shore. It may be that the high windexposure
and salt spray that occur on exposed headlands prevent the establishment
and growth of taller-growing shrubs.
The MDS analysis suggests there are regional differences in coastal heathlands
in Nova Scotia. Oberndorfer and Lundholm (2009) also suggest that Nova Scotia
coastal heathlands do not have repeating vegetation communities across regions.
Thus, each region within the province supports a unique assemblage of species in its
coastal heathlands. Conservation planning must consider these regional differences
in order to capture the diversity of community types found in coastal heathlands.
Protected or conserved heathlands in one part or region of the province may not
necessarily capture the diversity found in other areas of the province.
Species richness
Heathlands in Nova Scotia may be more species-rich than earlier studies
indicate. Davis and Browne (1996) suggested that heathlands in Nova Scotia
are nutrient-deficient, with low floral diversity and a small number of niches.
Strang (1971) reported low plant diversity for heathlands in the Tobeatic area.
In contrast, we found 102 species of vascular plants and 37 species of lichens
and mosses. Oberndorfer and Lundholm (2009) reported 173 species of vascular
plants, mosses, and lichens for their study in coastal heathlands in Nova Scotia.
The plant species richness found in heathlands is comparable to the richness
found in forests in Nova Scotia. Neily et al. (2011) reported 30–80 species of
vascular plants, mosses, and lichens in different forest vegetation types in Nova
Scotia. Moola and Vasseur (2004) reported only 53 species of ground plants in
their study of Picea rubens Sarg. (Red Spruce) communities in coastal Nova
Scotia. Glaser (1992) reported 81 species of vascular plants for raised bogs in
Nova Scotia. Cameron (2009) found 78 species of vascular plants and 35 species
of lichens and mosses in his plots in Acer rubrum (Red Maple) wetlands in Nova
Scotia. Heathlands in Nova Scotia are clearly comparable to other ecosystems
types and can no longer be considered as having low plant diversity.
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Rare species
Heathland rare plants are not restricted to any particular community type, but
occur in a wide variety of habitats. For example Alpine Whortleberry can be found
on exposed rock with xerophilic lichens in very dry habitats. Michaux’s Dwarf
Birch occurs in imperfectly- to poorly-drained habitat with extensive cover by
Sphagnum species. Furthermore, both the exposed-rock lichen community and the
imperfectly-drained Sphagnum community can be found in other coastal heathlands
without the associated rare species. Also, several rare species such as Alpine
Whortleberry and Michaux’s Dwarf Birch occur in both low- and high-shrub communities.
Oberndorfer and Lundholm (2009) also found rare species occurring in
a variety of habitats in their plots in Nova Scotia coastal barrens. These findings
mean that efforts toward conservation of rare coastal-barren plants will need to be
site-specific rather than aimed at certain community types.
We found that rare plants occurred in both low- and high-shrub communities;
however, more rare species occurred in low-shrub communities. Oberndorfer and
Lundholm (2009) found rare species richness was greater where vegetation height
was low. Rare plants in coastal heathlands in the northeastern US are mostly reported
as occurring in low-shrub communities (Dunwiddie 1990, Godfrey and Alpert
1985, Noss et al. 1995). We speculate that it is likely that some rare arctic-alpine
plants cannot compete with taller ericaceous shrubs found in high-shrub communities,
but are able to survive harsh conditions found on headlands where high-shrub
species cannot survive. However, coastal high-shrub communities should not be
overlooked in conservation planning, as they contain rare species, and also add to
the ecosystem diversity of terrestrial coastal systems.
Conclusion
Coastal heathlands are complex habitats, with community types and species not
found elsewhere in the province. Species richness is also high in these communities
compared to some forest and wetland communities in Nova Scotia, and rare species
occurrences are not necessarily correlated with specific community types. Both
high- and low-shrub coastal heathlands add diversity to the predominately-forested
landscape of the province and should be of conservation concern.
Given the various threats to these ecosystems observed during the course of this
study, conservation concern is especially warranted. All-terrain vehicles (ATV)
trails were numerous in the heathlands, even in protected areas where ATV are
prohibited. Plants were trampled and killed by vehicles where trails occur, and soil
erosion was evident on hills and slopes. Coastal development may also be a concern
for conservation of these ecosystems. Housing and cottage developments in coastal
high-shrub communities on private land were noted during the study. Development
not only can result in destruction of coastal heathlands, but may increase human
recreational-activity impacts as well.
Acknowledgments
We would like to thank those who helped with the field-work: Leif Helmer, Will Marten,
Dave Williams, Ron Williams, and Tammy Wilson. Thanks to Julie Towers for a helpful
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review of the manuscript and John LeDuc for supporting the work. We would also like to
thank Sean Blaney, Dr. Jeffrey Corben, and an anonymous reviewer for their helpful reviews.
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Appendix 1. Mean cover class values for plant species by occurrence, physiognomy, geographic
location, and layer for 39 plots in Nova Scotia heathlands. “-” indicates no occurrence.
n = number of plots; high = high-shrub, low = low-shrub.
Inland Coastal
Species high low low high
n 7 2 20 10
Canopy
Abies balsamea (L.) Mill. - - - 4
Acer rubrum L. 3 - - -
Larix laricina (Du Roi) K. Koch - - 1 2
Picea glauca (Moench) Voss - - 1 -
Picea mariana (Mill.) Britton, Sterns & Poggenb. 3 1 2 4
Pinus banksiana Lamb. - - 3 -
Pinus resinosa Aiton 2 - - -
Pinus strobus L. 3 1 - -
Subcanopy
Larix laricina - - 1 -
Picea mariana - - 2 -
Pinus banksiana - - 2 -
Shrub
Abies balsamea - - - 1
Acer rubrum - - 2 2
Alnus incana (L.) Moench - - - 2
Alnus viridis (Chaix) DC. 3 - 2 3
Amelanchier laevis Wiegand - - - 2
Amelanchier spp. Medik. 1 - 2 1
Andromeda polifolia L. - - 1 -
Betula papyrifera Marshall var. cordifolia (Regel) Fernald - - - 1
Betula michauxii Sarg. - - 1 5
Chamaedaphne calyculata (L.) Moench 4 - 2 3
Gaylussacia baccata (Wangenh.) K. Koch 6 4 4 3
Ilex glabra (L.) A. Gray 3 4 - 5
Ilex mucronata (L.) Powell, Savolainen & Andrews 4 3 1 3
Juniperus communis L. - 3 3 1
Kalmia angustifolia L. 3 4 2 4
Kalmia polifolia Wangenh - - 1 -
Larix laricina - - 1 -
Ledum groenlandicum Nutt. 1 - 2 2
Myrica gale L. 2 - 2 3
Morella pensylvanica (Mirb.) Kartesz 3 4 3 3
Osmunda cinnamomea L. - - 5 -
Photinia melanocarpa (Michx.) K.R. Robertson & Phipps 3 - 1 2
Picea mariana - - 1 3
Rhododendron canadense (L.) Torr. 3 3 - 3
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Inland Coastal
Species high low low high
Rubus hispidus L. 2 - - -
Sorbus Americana Marshall - - - 1
Spiraea alba Du Roi - - 1 -
Vaccinium angustifolium Aiton 2 1 2 3
Vaccinium boreale I.V. Hall & Aalders - - 2 1
Vaccinium uliginosum L. - - 3 2
Viburnum nudum L. 3 1 - 2
Herb
Abies balsamea - - - 2
Acer rubrum - - - 1
Amelanchier spp. - - 1 -
Angelica lucida L. - - 3 -
Aralia nudicaulis L. 2 - 4 1
Arctostaphylos uva-ursa (L.) Spreng. - 4 2 -
Aster sp. - - 4 -
Calamagrostis canadensis (Michx.) P. Beauv. - - 1 -
Carex nigra (L.) Reichard - - 1 -
Carex spp. 3 - 1 -
Carex trisperma Dewey - - 5 -
Chamaedaphne calyculata -` - 5 -
Clintonia borealis (Aiton) Raf. - - 3 1
Coptis trifolia (L.) Salisb. - - 1 2
Corema conradii (Torr.) Torr. ex Loudon - 5 4 2
Cornus canadensis L. 3 - 2 3
Dalibarda repens L. - - - 1
Deschampsia flexuosa (L.) Trin. - - 5 -
Drosera rotundifolia L. - - 1 2
Eleocharis palustris (L.) Roem. & Schult. - - 3 -
Empetrum eamesii Fernald & Wiegand - - 6 -
Empetrum nigrum L. - - 4 3
Epigaea repens L. - - 2 -
Eriophorum vaginatum L. - - 2 2
Gaultheria hispidula (L.) Muhl. ex Bigelow - - - 2
Gaultheria procumbens L. 3 3 2 2
Gaylussacia baccata - - 4 -
Iris prismatica Pursh ex Ker Gawl. - - 2 -
Iris versicolor L. - - 2 -
Juncus effusus L. - - - 2
Juniperus communis - - 3 -
Juniperus horizontalis Moench - - 3 -
Kalmia angustifolia - - 3 5
Kalmia polifolia - - 2 -
Ledum groenlandicum - - 2 -
Lonicera caerulea L. - - - 2
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2013 Northeastern Naturalist Vol. 20, No. 4
Inland Coastal
Species high low low high
Lycopodium clavatum L. - - 1 -
Lycopodium obscurum L. - 1 1 -
Maianthemum canadense Desf. 1 - 2 2
Medeola virginiana L. - - 1 -
Melampyrum lineare Desr. - - 1 2
Minuartia glabra (Michx.) Mattf. - - 2 -
Myrica gale - - 2 -
Moehringia lateriflora (L.) Fenzl - - 1 -
Morella pensylvanica - - 2 -
Osmunda cinnamomea - - - 1
Photinia melanocarpa - - 2 2
Picea glauca - - 2 1
Picea mariana - 4 2 -
Pinus banksiana - - 1 -
Pinus strobus - - - 1
Plantago maritima L. - - 2 -
Prenanthes trifoliata (Bigelow) Torr. - - 2 1
Pteridium aquilinum (L.) Kuhn 5 4 2 4
Pyrola elliptica Nutt. - - - 1
Ribes hirtellum Michx. - - 1 -
Rosa nitida Willd. - - 5 -
Rubus chamaemorus L. - - 2 2
Rubus pubescens Raf. - - 2 2
Sanguisorba canadensis - - 2 -
Sarracenia purpurea L. - - 2 2
Sibbaldiopsis tridentata (Aiton) Rydb. - - 2 -
Smilacina stellata (L.) Link - - - 1
Smilacina trifolia (L.) Sloboda - - 1 -
Solidago sempervirens L. - - 2 -
Thalictrum pubescens Pursh - - 1 -
Trichophorum cespitosum (L.) Hartm. - - 3 -
Trientalis borealis Raf. - - 1 1
Vaccinium angustifolium - - 2 4
Vaccinium macrocarpon Aiton - - - 1
Vaccinium oxycoccus L. - - 2 4
Vaccinium vitis-idaea L. - - 1 2
Viburnum nudum L. - - 1 -
Viola spp. - - 2 -
Moss
Andreaea rupestris Hedw. - - 3 -
Dibaeis baeomyces - - 3 -
Bazzania trilobata (L.) A. Gray - - 1 1
Bryum argenteum Hedw. - - 1 -
Cetraria islandica (L.) Ach. - - - 2
Cetaria muricata (Ach.) Eckfeldt - - - 3
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Inland Coastal
Species high low low high
Cladonia arbuscula (Wallr.) Flotow 3 4 3 -
Cladonia boryi Tuck. - - 2 -
Cladonia maxima (Asahina) Ahti - - 1 -
Cladonia pleurota (Flörke) Schaerer - - 2 -
Cladonia rangiferina (L.) F.H. Wigg. - 4 2 2
Cladonia scabriuscula (Delise) Nyl. - - - 1
Cladonia spp. - - - 3
Cladonia stellaris (Opiz) Pouzar & Vězda - - 2 2
Cladonia stygia (Fr.) Ruoss 3 4 - -
Cladonia terrae-novae Ahti - - 3 3
Dicranum majus Sm. - - - 3
Dicranum polysetum Sw. - - 2 1
Dicranum scoparium Hedw. - - 1 1
Dicranum spp. 2 1 1 -
Dicranum undulatum Sw. - - 2 1
Hylocomium splendens (Hedw.) Schimp. 2 - 2 1
Hypnum spp. - - - 1
Leucobryum glaucum (Hedw.) Ångstr. - - 2 -
Parmelia saxatilis - - 1 -
Pleurozium schreberi (Brid.) Mitt. 3 - 3 2
Polytrichum commune Hedw. - - 2 2
Polytrichum juniperinum Hedw. - - 3 -
Sphaerophorus fragilis (L.) Pers. - - 1 -
Sphagnum capillifolium (Ehrh.) Hedw. - - - 3
Sphagnum compactum DC. - - 5 -
Sphagnum fallax (Klinggr.) Klinggr. 3 - - 5
Sphagnum fuscum (Schimp.) Klinggr. - - 3 3
Sphagnum magellanicum Brid. - - 1 5
Sphagnum papillosum Lindb. - - - 4
Sphagnum russowii Warnst. - - 2 -
Sphagnum spp. 4 3 4 1
Sphagnum warnstorfii Russow - - 1 -
Stereocaulon dactylophyllum Flörke - - 3 -
Stereocaulon spp. - - 2 -