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Research article

Effects of land ownership and landscape-level factors on rare-species

richness in natural areas of southern Ontario, Canada
J. Lovett-Doust
, M. Biernacki
, R. Page
, M. Chan
, R. Natgunarajah
and G. Timis
Department of Biological Sciences, University of Windsor, Windsor, Ontario, Canada N9B 3P4;
of Biology, University of Memphis, Memphis, TN, 38152-3540 USA;
Author for correspondence (e-mail:
Received 28 March 2002; accepted in revised form 6 May 2003
Key words: Biodiversity, Conservation, Habitat, Land ownership, Landscape-level effects, Land-use, Natural ar-
eas, Rare species fragmentation
Surprisingly few studies have considered the extent to which the nature of the ownership of land is associated
with differences in biodiversity. We analysed ownership and other landscape-level effects on rare-species rich-
ness for both globally- and regionally-rare biota including birds, herpetofauna, butteries, mammals, and plants
in 289 designated natural areas NAs in southern Ontario, Canada. Information about each NA including
area, number of plant communities, ownership status and details of species diversity were collected from pub-
lished sources. Length of perimeter of NA, relative isolation, and an estimate of fragmentation were measured
using image analysis and GIS techniques. NAs were in general relatively small, with mean area of 158 ha me-
dian 85 ha, range from 0.9 to 1278 ha for private NAs; public NAs had mean area of 132 ha median 16 ha,
range from 0.1 to 1481 ha. Mean number of plant communities was 4.6 median 4, range 1- 13 at private NAs
and 3.8 median 3, range 1-16 at public NAs. Our results show that, of several landscape-level factors, area had
the greatest effects on rare-species richness and other biotic indices. Effects of area were followed by effects of
plant community diversity, however this was itself signicantly affected by area and the extent of perimeter of
the NA. Both these factors were followed by effects of ownership of the NA and by effects of isolation of the
NA represented by minimum distance to nearest NA and by number of NAs in 10 km radius. Other landscape-
level factors did not appear to have overall signicant effects. Variation in area accounted for 0.1% to 29% of
variation in number of rare species, with lower values for globally-rare, than for regionally-rare taxa. For all
biotic groups, public ownership of NAs was associated with signicantly greater rare-species richness compared
to private ownership, even after other factors such as area were controlled. For all globally-rare biota except
butteries, area of NA had greater effects on rare-species richness than did ownership. Richness of regionally-
rare birds was more affected by plant community diversity than by area of NA. Number of recorded plant com-
munities accounted from 2.1% of variation in number of globally-rare plant species to as high as 31% of variation
in regionally-rare butteries. The diversity of plant communities was itself inuenced by total site area account-
ing for 45% of variation, extent of elongation of the NA, and both external- and interior- edge perimeters. Pub-
lic NAs had greatest numbers of rare biota and so should be a signicant focus for conservation programs.
Smaller, privately-owned patches of natural area dominate by number and area in this densely populated region
and their signicance should not be overlooked.
2003 Kluwer Academic Publishers. Printed in the Netherlands.
Landscape Ecology 18: 621633, 2003.
Land-use change and ecosystem exploitation have
been the primary forces causing loss of biological di-
versity Daily 1997; Vitousek et al. 1997; Sala et al.
2000. Land ownership patterns are obviously linked
to landscape alteration for agricultural, industrial and
residential needs, yet few studies have considered the
extent to which the nature of the ownership of the
land is associated with differences in biodiversity but
see Kindscher and Scott, 1997; Crow et al. 1999;
Lovett-Doust and Kuntz 2001. In southern Ontario,
a majority of the land exists under private ownership.
Much of this is used agriculturally, leaving tracts of
fragmented natural areas also present Pearce, 1993.
In the United States too, private land ownership
dominates nearly 67% of land in the contiguous
U.S. is privately held, another 2% is owned by state
or local governments, and the federal government
owns 263 million ha, or 31% Dale et al. 2000;
Groves et al. 2000. Developed, non-federal lands
there have increased by 18% in the past decade, to 92
million ha or 4.4% of the total area Dale et al. 2000.
In Ontario, projected population growth by 34% to
c. 16.6 million by 2028 will further increase pressure
on remaining natural areas Government of Ontario,
2000. Privately-owned lands, both globally and in
southern Ontario, have been under more intensive
economic pressure than public lands in satisfying hu-
man needs for goods and services Pearce 1993; Ri-
ley et al. 1996; Daily 1997; Vitousek et al. 1997.
Thus effects of land ownership on biodiversity in re-
maining natural areas need to be considered in con-
servation programs.
The extent of biodiversity maintained on private
versus public lands seems likely to be inuenced by
factors directly associated with the purpose of the
land ownership Lovett-Doust and Kuntz 2001. It
could also be affected by factors indirectly related to
ownership, such as the landscape characteristics of
private versus public lands. For example, if public
lands tend to be found in larger and less fragmented
habitat blocks, they could be expected to contain
more biodiversity even if the selection of lands, or
their management by private or public entities, was
identical see Groves et al. 2000. MacArthur and
Wilson 1967 used area as a single, major factor,
predicting species richness on oceanic islands. The
well-known species-area relationship is overwhelm-
ingly important in terrestrial ecosystems, however
patches may also be affected by other, landscape-level
factors in addition to size, such as the number of
landforms present and the extent of interior forest
habitat Lovett-Doust and Kuntz, 2001. Patterns of
biotic diversity at public versus private lands has ob-
vious pertinence to conservation and public planning,
because of both the direct and indirect effects noted.
Here we investigate patterns of rare and threatened
biodiversity including birds, herpetofauna, butter-
ies, mammals, and plants across land ownership
categories, in landscape patches containing largely
natural vegetation . We also examined the extent to
which differences may be explained by indirect,
landscape-level features including the length of the
patch perimeter, the patch perimeter-to-area ratio, and
the relative isolation of the site from other natural ar-
eas. We analysed the numbers of both globally-, and
regionally-rare biota, from an array of natural areas
in southern Ontario, as indices of regional biodiver-
sity, in order to evaluate the conservation importance
of land ownership and these other, landscape-level
Study region
We studied the distribution of globally- and region-
ally-rare organisms in 344 protected natural area
NAs; see following section for details in southern
Ontario, Canada see Figure 1. This region has been
exposed to intense anthropogenic pressure, possibly
greater than any other region in Canada. Geographi-
cally, the region comprises less than one percent of
Canadas land area yet contains 25% of the popula-
tion Government of Ontario 2000, with some esti-
mates projecting 40% by 2026 Government of
Ontario 2000. Southern Ontario sits at the northern
edge of the Eastern Deciduous Forest Region of
North America Theberge 1989; Delcourt and Del-
court 2000; natural vegetation includes deciduous
forest, tallgrass prairie and savanna, and wetland sys-
tems. There is considerable climatic amelioration
from the Great Lakes Theberge 1989, which has
favoured European settlement and agriculture over
the past three centuries. Native people used the region
agriculturally, prior to European settlement. Develop-
ment of the region was also facilitated by the ready
supply of rock and aggregate in the Niagara Escarp-
ment, a 750 km, largely forested, Paleozoic limestone
escarpment that is the most prominent topographical
feature in the region, after the Great Lakes Riley et
al. 1996; Lovett-Doust and Kuntz 2001. We have
previously demonstrated the importance at natural ar-
eas of the Niagara Escarpment World Biosphere Re-
serve of land ownership, as well as certain landscape-
level factors associated with the escarpment
environment and not examined here including the
number of geological landforms, and the extent of in-
terior forest on various biotic indicators Lovett-
Doust and Kuntz 2001. We also recently reported
signicant effects due to land-use in the background
matrix of land surrounding these escarpment sites
Biernacki M., Lovet-Doust J., Page R.; unpublished
Historically-continuous deciduous forest in the re-
gion has decreased signicantly and become highly
fragmented Pearce 1993; Larson et al. 1999. At
present, forest cover ranges from 7 40% mean
24.8% of its pre-settlement extent Larson et al.
1999. Consequently remaining species have been
concentrated within smaller, more isolated, natural
areas that exist scattered across the regional land-
Data collection
An electronic database was created using published
information for physical and biological features at
344 remnant areas of natural habitat, all of which had
previously been designated as environmentally signif-
icant areas ESAs by Ontario Conservation Authori-
ties Oldham 1983; Heagy 1993, 1995. These natural
areas had been identied as important and valuable
because: 1 they contained particular natural features
such as signicant geological landforms, high levels
of biodiversity, rare species, or rare community
types; 2 they provide important services for wildlife
e.g., serving as migratory staging grounds, or part of
Figure 1. Map of study area encompassed by thick line with 289 NAs in southern Ontario, Canada. The Niagara Escarpment Biosphere
Reserve is marked by shaded area. Lines indicate shorelines of lakes and rivers.
a habitat linkage system; or 3 they offer particular
value for aesthetic, recreational, scientic, or educa-
tional purposes. The designation of environmentally
signicant confers a certain level of recognition
upon a NA, but otherwise carries no particular pro-
tective status Oldham 1983; Heagy 1993, 1995.
Biotic inventories mostly published within the
past 10-12 years of NAs had been carried out by ex-
perienced biologists from either the Ontario Ministry
of Natural Resources OMNR or regional Conserva-
tion Authorities, and utilized extensive eld studies,
faunal trapping, and surveys Table 1. The primary
purpose of these inventories had been to characterize
regional features as fully as possible, hence intensity
and accuracy of eld surveys were similar across
natural areas, regardless of ownership status. Our da-
tabase was developed using published results of in-
ventories for 344 NAs, though complete information
about both area and ownership were available for
only 289 NAs. Table 1 summarizes data sources for
these NAs.
Species rarity ranks
We determined regional distribution and abundance
status for each species reported as present at a NAand
designated as having conservation signicance in
Ontario i.e., identifed as being rare, in terms of ei-
ther its global distribution and abundance G-rank
or its regional distribution and abundance S-rank.
G- and S-ranks have ve point scales, with 1 repre-
senting the very rarest at either global or regional
scales, and 5 representing the most common Master
1991. Information was determined for mammal, am-
phibian, reptile, bird and buttery faunal groups from
the Natural Heritage Information Centre, Ontario
Ministry of Natural Resources, Peterborough, Ontario
NHIC 1999. Global and regional rarity ranks for
plants were taken from Newmaster et al. 1998. Fol-
lowing convention, in our analyses species designated
as either G1, G2, or G3 were regarded as globally
rare and species designated as S1, S2, or S3 were
classed as regionally rare. For taxa with uncertain
conservation status e.g., S3S4, or G1G2, the taxon
was conservatively assigned the more rare designa-
tion S3 or G1, respectively, in the above example.
Land ownership
Ownership of each NA was assessed from individual
site descriptions in the ESA reports, and NAs were
assigned to one of four ownership categories: private;
public; mixed ownership; First Nations tribal/band
owned. There were so few tribally-owned NAs that
this category was excluded in analyses. Private own-
ership meant the NA was owned and managed by ei-
ther one or more private landowners. In contrast a
publicly-owned NA was owned and managed either
by some levels of government municipal, county,
provincial or federal, or by a public agency, such as
an Ontario regional Conservation Authority. In many
cases, part of the area of a NA belonged to a public
jurisdiction while the reminder was privately owned;
these were treated as mixed ownership.
Landscape parameters of NAs
Digitized maps for each NA available from Ontario
Ministry of Natural Resources OMNR, or regional
Conservation Authorities were used to calculate spa-
tial parameters, including area, perimeter and an es-
timate of shape. Digitized maps of the same scale
1:25,000 were analysed using ArcView software v.
3.2 Environmental Systems Research Institute, Red-
land, CA. For each NA, we determined area and pe-
rimeter relations as follows.
Table 1. Data sources for natural areas in southern Ontario, by locality.
Locality Number of NAs Data sources
total analysed
Essex County 50 50 Environmentally Signicant Area ESA data assembled using information from Oldham
1983 updated 1991 and Kamstra et al. 1995
Hamilton-Wentworth 102 85 Heagy 1993, 1995
Metropolitan Toronto 135 117 Metropolitan Toronto and Region Conservation Authority 1982, plus updates of 1994,
1995, and 1996
Lower Trent Region 38 37 Brownell and Blaney 1995
NAs for which data about area and ownership were available
An index of perimeter-to-area was calculated to re-
present the degree of fragmentation for each NA. We
used the sum of the external perimeter P
plus the
cumulative length of interior perimeter P
ated by any roads, railroads or powerline corridors
within an NA:
Perimeter-to-area ratio


where A area of NA. Isolation of NAs was evalu-
ated as the shortest distance from the perimeter of one
NA to the perimeter of the next-nearest NA. The
number of other NAs within a 10 km contour, radiat-
ing from the patch perimeter, was determined for each
Estimates of habitat diversity were made for each
NA following the Ecological Land Classication for
Southern Ontario Lee et al. 1998. This system rec-
ognizes six nested levels, ranging from site region,
through community series, to vegetation type. We de-
termined the number of community series in each NA
e.g., swamp, marsh, deciduous forest, grassland.
Statistical analyses
Effects of landscape-level traits of NAs on numbers
of rare species per NA were analysed using analysis
of variance, analysis of covariance, and linear regres-
sion using Systat Systat for Windows v.10, SPSS Inc.
Chicago, IL. Ownership was treated as a categorical
variable; area, perimeter length, perimeter-to-area ra-
tio, extent of elongation longest axis tted in the
contour of NAdivided by longest perpendicular axis,
and isolation distance to nearest NA were treated as
continuous variables. In multi-factorial regressions, a
standard stepwise backward procedure was used to
identify the most signicant independent variables
and to maximize power and determination of the co-
efficient of regression Zar 1999; Systat for Windows
v.10, SPSS Inc. Chicago, IL. Pearson correlations of
species richness of rare biota and associated probabil-
ities were also evaluated. A non-linear procedure was
used to estimate the optimal exponent of power-based
species-area relationship functions for each group of
biota according to the following model:
S number of species in biotic group;
C coefficient for area;
A area of NA;
Z exponent for area.
A least squares procedure utilizing the Gauss-
Newton algorithm and available in Systat was used
to estimate the optimal parameters of species-area re-
lationship. Prior to linear analyses, all variables were
evaluated for normality of distribution and, when
necessary, log n1 transformations were applied
Zar 1999.
The array of rare species represented in individual
NAs was quite low in terms of absolute numbers
within biotic groups present in any one NA, however
there was considerable heterogeneity Table 2. For
example, at several NAs, all three of Ontarios glo-
bally rare herptile species occurred; at another site, 49
bird species, or 68% of all Ontarios regionally-rare
bird species were recorded. Indeed, two of a total of
eleven and 49 of 72 globally-rare, and regionally-
rare bird species, respectively, occurred in single
NAs; yet overall there were just two NAs which con-
tained more than twenty, regionally-rare birds. Nearly
all NAs having any regionally-rare birds at all, con-
tained just 1 4 of these species. Regionally-rare
plants with 732 species recorded in Ontario had the
greatest general representation, yet on average only c.
6% of all regionally-rare plant species occurred at
each NA.
About half the NAs 157 sites were owned
completely by private interests, a further quarter 77
sites by a mixture of private and public ownership,
while about a fth 55 sites were owned by one or
more public jurisdictions. The frequency distribution
of sizes, and the mean area of NAs in each owner-
ship group differed Figure 2. Privately-owned NAs
dominated; by area, privately-owned NAs contributed
nearly 49% of the total area of analysed NAs, while
mixed ownership gave 36%, and public ownership
just 15%.
Analysis of covariance with area treated as cova-
riate identied ownership as a signicant inuence
on rare-species richness. With the exception of glo-
bally-rare birds and herptiles, and regionally-rare
mammals, ownership had signicant effects on the
numbers of all groups of rare species Table 3. Fur-
thermore, wherever there was a signicant difference,
publicly-owned sites had signicantly greater num-
bers of rare biota than other forms of ownership
Table 3. For globally-rare butteries, ownership had
even greater effects than did effects of area. In the
case of globally-rare butteries, ownership accounted
for over 12% of variation, while area alone explained
only 0.15% of variation see Table 3.
Area per site ranged from 1 ha to 2400 ha Figure
2. Most sites were relatively small; more than three
quarters of NAs had area 100 ha and more than
half the NAs had area 65 ha. Mean and median ar-
eas were 163.3 ha SE 17.1 and 64.6 ha, respec-
tively. Only three sites had area 2000 ha. For
private sites, mean area was158 ha median 85 ha,
range 0.9 to 1278 ha, and for public sites it was132
ha median 16 ha, range 0.1 to 1481 ha. At mixed
ownership sites the mean was 249 ha median 61 ha,
range 0.5 2400 ha. A nonparametric Kruskal-
Wallis test indicated that private NAs were signi-
cantly larger than mixed and public sites test
statistic11.63, P0.009, df2. Areas of public and
mixed ownership did not differ signicantly in size.
Analysis of covariance showed that without regard
to type of organism, 28.3% of variation in the num-
ber of rare species was explained by variation in area.
Overall, variation in area accounted for 0.2% to 29%
of variation in rare species numbers, with lower val-
ues for globally-rare taxa than for regionally-rare
ones Table 3. In general, with increasing area there
was a signicant R
0.04; df1, 219; F9.13;
p0.003 decrease in distance to nearest NA; there
was also a signicant relationship R
0.021; df1,
221; F4.80; p0.03 between size of NA and num-
ber of other NAs within a 10 km distance Figure 3.
Smaller sites were more isolated and were less likely
to have other NAs within a 10 km distance. Isolation
or number of other NAs varied enormously for small
NAs from extremely isolated to not isolated at all,
whereas degree of isolation for large NAs was con-
sistently low Figure 4
Landscape traits of NAs
Numbers of species of both globally- and regionally-
rare biota decreased signicantly R
0.43; df1,
44; F29.75; p 0.001 and R
0.34; df1, 203;
F104.1; p 0.001 respectively for globally- and
regionally-rare biota per unit area as the size of NA
increased Figure 4. The smallest NAs were most
likely to have the greatest number of rare biota per
unit area. Shorter distance to other NAs was signi-
cantly R
0.07; df1, 203; F6.61; p0.011 re-
lated to larger numbers of rare biota at a NA. In
contrast to effects of area, perimeter-to-area ratio did
not have signicant direct effects on the number of
rare species at a NA. However, regression analyses
indicated a highly signicant R
0.18; df1, 202;
F19.84; p 0.001 negative relationship between
perimeter-to-area ratio and NA plant community di-
Table 2. Distribution patterns for globally- and regionally-rare biota in natural areas NAs in southern Ontario. N number of NAs having
data for a particular biotic group.
Biotic Group N Total number of
species re-
corded in On-
Maximum num-
ber of species
present in an
individual NA
% of total
Mean number
SE of spe-
cies at each NA
Number of NAs having particular num-
bers of rare species:
0 1-4 5-10 11-20 20
Birds Globally rare 240 11 2 18% 0.02 0.01 238 2 0 0
Regionally rare 278 72 49 68% 0.76 0.24 210 63 3 0 2
Butteries Globally rare 94 5 2 40% 0.15 0.08 90 4 0
Regionally rare 94 44 14 32% 0.78 0.19 60 32 1 1 0
Mammals Globally rare 232 1 0 0 0 0
Regionally rare 232 15 5 33% 0.06 0.03 223 8 1 0
Herptiles Globally rare 168 3 3 100% 0.18 0.04 150 18
Regionally rare 236 18 12 66% 0.34 0.08 204 27 4 1
Globally rare 242 32 6 19% 0.14 0.04 222 19 1 0 0
Regionally rare 289 732 113 15% 4.70 0.62 157 85 29 26 19
versity. Furthermore plant community diversity had
highly signicant effects on rare-species richness in
all biotic groups, except for globally-rare butteries
Table 4.
The number of other NAs in a 10 km radius from a
NA was signicantly correlated with number of rare
species observed R
0.07; df1, 191; F6.79;
p 0.008. Effects of the presence of other NAs
within a 10 km radius were signicant for richness of
globally-rare biota R
0.04; df1, 44; F4.97; p
0.031 and for regionally-rare biota R
0.12; df1,
201; F9.62; p 0.002.
Biotic variables
Mean number of plant communities was 4.6 median
4, range 1- 13 at private NAs, 4.1 median 3, range
1-12 at mixed ownership NAs, and 3.8 median 3,
range 1-16 at public NAs. The nonparametric
Kruskal-Wallis test did not indicate signicant differ-
ences in plant community diversity among the three
ownership groups test statistic4.24, P0.24,
df2. Univariate regression analyses indicated that
community diversity was highly signicantly corre-
lated with the number of rare biota at each NA. Re-
gression coefficients differed for each group of rare
biota and were greatest for regionally-rare plants.
Community diversity accounted for between 2.1% of
variation in species number for globally-rare plants
to 31% of variation in species number for regionally-
rare butteries. Furthermore, for all but regionally-
rare birds, the proportion of variance in values for
rare animal species richness accounted for by com-
munity diversity exceeded the variance accounted for
by area e.g., 2.8% vs 13.9% for globally-rare birds;
Table 4. In contrast, variance in the number of rare
plant species was better accounted for by the area of
NAs than by community diversity. Area was highly
signicant N289, p 0.001 and itself accounted
for c. 45 % of variation in plant community diversity.
Perimeter-to-area ratio also had highly signicant ef-
fects N289, p 0.001 and accounted for a further
3% of variation in community diversity. Thus, both
area and the perimeter-to-area ratio had possible in-
direct inuences on rare biota suggested by their cor-
relation with community diversity.
Figure 2. Frequency distribution of area of NAs in different own-
ership classes.
Public NAs in Ontario harbour signicantly greater
numbers of species, particularly of globally and re-
gionally rare biota, than do private NAs. This obser-
vation in particularly striking because, in southern
Ontario, private NAs dominate by number, by cumu-
lative area and by average area of NA. This situation
may be a consequence of historical events related to
protection of particular NAs, and implementation of
conservation programs at the NAs. Yet despite a ma-
jor importance of public NAs in protecting rare biota
in southern Ontario, both public and private NAs har-
bour unique rare species found nowhere else, thus
complementing each other in conservation of rare
biota Heagy 1993, 1995. In contrast, preliminary
studies of rare-species diversity in the United States
suggest that private lands harbour rather greater num-
bers of rare and imperiled species than public lands
USGAO 1995; Stein et al. 1995; Groves et al. 2000.
Our results show that, of several landscape-level
factors, area had the overall greatest effects on rare-
Table 3. Summary of results of ANCOVA showing effects of NA ownership on numbers of globally and regionally rare biota. Area as
exponential function of NA was a covariate in all models. Each ownership category was ranked numerically in decreasing order, based on
rare species frequencies different lowercase letters indicate signicant ownership differences. Signicance is indicated as: * p 0.05; **
p 0.01; *** p 0.001.
Biota N R
F- ratio for ownership Ownership rank Exponent for area F- ratio for area
Public Mixed Private
Globally rare birds 237 0.039 1.157 1a 2a 3a 0.009** 8.196**
Regionally rare birds 274 0.110 8.604*** 1a 2b 3b 0.110*** 23.187***
Globally rare butteries 117 0.125 7.977*** 1a 2b 3b 0.004 0.074
Regionally rare butteries 117 0.257 3.287* 1a 3b 2ab 0.220*** 30.951***
Regionally rare mammals 237 0.025 1.758 1a 3a 2a 0.006 3.411
Globally rare herptiles 237 0.023 1.577 1a 2a 3a 0.018* 5.272*
Regionally rare herptiles 237 0.279 2.980* 1a 2ab 3b 0.057*** 16.440***
Globally rare plants 289 0.050 3.313* 1a 2ab 3b 0.022* 8.422**
Regionally rare plants 289 0.100 5.580*** 1a 2b 3b 0.203*** 21.867***
Globally rare biota 289 0.065 4.071** 1a 2b 3b 0.037** 13.363***
Regionally rare biota 289 0.121 5.174** 1a 2b 3b 0.270*** 32.645***
Total rare biota 266 0.283 2.952* 1a 2ab 3b 0.618*** 92.645***
Figure 3. Relationship between area of NA and A distance to nearest NA; and B number of NAs within a 10km distance from perimeter
of NA. Lines represent linear regression and 95% condence interval.
species richness and other biotic indices. Effects of
area were followed by effects of plant community di-
versity, however this was itself signicantly affected
by area and the extent of perimeter a function of
shape of the NA. Both these factors were followed
by effects of ownership of the NA, and by effects of
isolation of the NA from other NAs represented by
minimum distance to nearest NA, and by number of
NAs in 10 km radius. Other landscape-level factors
did not appear to have overall signicant effects.
Ownership of land
The concept of private ownership of land is one of
the most important structural attributes of North
American society, and it imparts great leeway to the
owner in land-use decisions. These directly impact
biota utilizing the land as habitat, migratory staging
points, etc. Platt 1996; Vitousek et al. 1997; Daily
1997; Dale et al. 2000; Government of Ontario,
Land-owners experience economic pressure to
convert land-use to development, or for agricultural
products Costanza et al. 1997; Adger and Luttrell
Figure 4. Number of A globally-rare biota-, and B regionally-rare biota per unit area ha of NA.
Table 4. Proportion of variation in rare species richness accounted for by NA area and plant community diver-
sity. Signicance is indicated as: * p 0.05; ** p 0.01; *** p 0.001
Biotic group Coefficients of determination R
Area Community diversity
Birds Globally rare 0.028** 0.139***
Regionally rare 0.118*** 0.106***
Butteries Globally rare 0.002 0.022
Regionally rare 0.288*** 0.312***
Mammals Globally rare
Regionally rare 0.005 0.065***
Herptiles Globally rare 0.017* 0.075***
Regionally rare 0.070*** 0.166***
Vascular plants Globally rare 0.019* 0.021**
Regionally rare 0.096*** 0.056***
2000. Lack of clear understanding of the values of
biodiversity and related ecosystem services and, in-
deed, inadequate bases for quantifying these values,
means these are often given too little weight in deci-
sions about land conversion Costanza et al. 1997.
Three studies have mapped the overall distribution
of species and land ownership in the U.S. Stein et al.
1995 examined the importance of U.S. federal lands
for maintaining federally-listed, endangered species:
only half of such species occurred at all on federal
lands. A study by the U.S. Government Accounting
Office USGAO 1995 reached a similar conclusion
through different methodology: one-half to two-
thirds of federally-listed species occurred non-exclu-
sively on federal lands. Finally, Groves et al. 2000
examined the distribution in the U.S. of species with
federal status, and assessed the distribution of those
species classied as globally imperiled i.e., having
G1 or G2 status. Federal land supported nearly 60%
of the federally-listed endangered species and about
the same fraction of imperiled species. State land
contained at least one population for 43% of imper-
iled species and 58% of federally-listed species.
Among major federal agencies, Dept. of Defense
lands appeared to contain the greatest numbers of
federally-listed species, while Forest Service lands
had most imperiled species. In contrast, private land
in the U.S. holds at least one population of 50% of
all imperiled species and c. 66% of federally-listed
species Groves et al. 2000. Southern Ontario has
one of the greatest fractions of land in private own-
ership in Canada Ramsey 1988; Government of On-
tario 1997. Private NAs dominate numerically and
by area about three times the number and the area of
public NAs see Figure1. Yet numbers of rare biota
and plant communities per unit area and per NA were
greatest on public lands Table 3. Our results corre-
spond to the general conclusions of Thomas et al.
1997 who determined that old woodlands 400
years in Britain had greater species richness when on
publicly- and trust-owned lands than when they were
located on private- and estate-owned lands.
Spies et al. 1994 found that over the course of 16
years, initially-continuous conifer forests on public
lands had three times more area overall, with more
closed canopy and three times more interior forest,
than otherwise comparable, privately-owned lands. In
southern Ontario, studies have shown that the largest
intact patches of forest were publicly owned, under
the jurisdiction of Ontario Ministry of Natural
Resources Pearce 1993. Furthermore, privately-
owned lands surrounding nearby public lands were
more fragmented and had highly isolated woodlots
Pearce 1993. Similarly, in an analysis of 98 envi-
ronmentally signicant natural areas along the Nia-
gara Escarpment in southern Ontario, Lovett-Doust
and Kuntz 2001 found that NAs with the largest
area and greatest extent of forest-interior habitat were
under mixed private and public ownership. Private
and public NAs there did not differ in mean size, yet
after controlling for the area of the site, and for the
extent of interior habitat and the degree of landform
heterogeneity, biotic diversity remained signicantly
greater on public land than on private Lovett-Doust
and Kuntz 2001.
We recognize obvious limitations to our database
and study. For example, ownership was determined at
a single occasion, at the time of the NAs inventory,
and there was no record of ownership over time. It
seems likely that some of the best lands in southern
Ontario suitable for parks i.e., large and highly
biodiverse may have been purchased in the past for
public ownership. Parks are often selected because of
their excellent representation of regional features, and
rare-species richness. Other NAs may have been
publicly acquired more recently, or may have been
transferred as donations from private or mixed own-
ership. Furthermore, simple presence of rare species
at a location says nothing about the population via-
bility or quality of the NA for rare species. Perhaps
most importantly, we have no information about the
management practices applied at NAs over time. Fi-
nally we note that, historically, land privatization did
not occur randomly, with respect to habitat types.
Stressful environments have tended to be the last that
were privatized, while fertile lands were rst. Gobin
et al. 2001 studied patterns of spread of land own-
ership in Nigeria, and observed that lands around
settlements and markets were most desired, followed
by land around roads and waterways, and this general
pattern spread as the human population grew and new
settlements and markets appeared. Lands far from
markets, roads, waterways, with unsuitable landforms
were least desired and remained under communal
ownership Gobin et al. 2001. Preferences of
particular lands for private ownership have been
noted elsewhere in North America, with more
publicly-owned lands occurring at higher elevation
and on soils less suitable for agricultural production
Spies et al. 1994; Craig et al. 2000; Wright et al.
2001. These lands were less desired for private own-
ership because of lower suitability for agriculture and
forestry, and difficulties with access Spies et al.
1994; Craig et al. 2000; Wright et al. 2001. Yet it
should be noted that low-fertility soils neednt neces-
sarily mean low biodiversity. Among the most biodi-
verse sites in the western United States are serpentine
outcrops in California with a tremendous number of
rare species, specically because of their low fertility
Ricketts et al. 1999. Similar observations have been
reported for New Zealand Craig et al. 2000 where,
despite the fact that 30% of all land is in reserves,
most conservation land occurs in areas that are not
useful for production. The same conclusions seem
likely for the intensely settled and highly agricultur-
alized land of southern Ontario.
Government and other public sector investment is
clearly important for the conservation of biodiversity,
and for several reasons may even be preferable to
private sector investment see James et al. 2000. At
present, the private sector is less prepared to provide
public goods which are related to the global environ-
ment such as the values and services of biological
systems and resources Wilson 1992; James et al.
2000. Estimates of economic value for the goods and
services that undisturbed natural areas provide are
sorely needed to bolster any economic argument for
protection of natural areas and to counterbalance the
more readily apparent short-term benets of develop-
ment Cairns and Bidwell 1996; Costanza et al. 1997;
Daily 1997.
Landscape-level traits of NAs
Results of this study indicate that area was the most
important factor, accounting for about a third of total
variance in numbers of rare species. However, effects
of area differed among ownership classes for numbers
of both globally- and regionally-rare species, with
signicantly larger numbers on comparably-sized
public land than on private Table 3. From the per-
spective of rare-species conservation, greater biodi-
versity of rare species per unit area occurred in small
sites compared to larger ones Figure 4. Of course,
simple presence alone does not indicate that popula-
tions are viable, especially for animal species. Our
results show for some biota, plants in particular, that
even relatively small areas supported a disproportion-
ately large number of rare species though we note
that, at least for birds, problems affecting survivorship
and fecundity occur disproportionately often in
smaller woodlots Burke and Nol 2000. Further-
more, depending on location, numerous small NAs
may serve as temporary staging areas and transfer
corridors for biota that migrate among habitat patches
Falkner and Stohlgren 1997; Keitt et al. 1997; Hale
et al. 2001. While smaller NAs may be sufficient for
protection of rare plants, most animals amphibians,
reptiles, mammals and birds, may require substan-
tially larger tracks of land for their home ranges. Our
results indicate that NAs in southern Ontario were
generally too small to provide habitat for globally and
regionally rare mammals.
Few effects of fragmentation estimated as perim-
eter-to area ratio were observed; the number of plant
communities was the sole variable responsive to
fragmentation. However, fragmentation effects on
plant community diversity may have carry-over
effects on rare biota, since rare-species richness was
signicantly dependent on these communities. Burke
and Nol 2000 found woodlot area was the most im-
portant factor affecting reproductive success of
forest-breeding birds in Ontario, with local plant
community diversity having no apparent effects. By
comparison, Freemark and Merriam 1986 observed
signicant effects of plant community diversity,
greater than the effects of habitat area, on diversity of
bird species in fragmented forests of southern
Ontario. Plant community itself was found to be sig-
nicantly affected by perimeter-to-area ratio. In sum,
results of our study suggest that area was the domi-
nant factor affecting species richness in NAs,
followed by effects of plant community diversity,
ownership, perimeter-to-area ratio, and isolation.
We are grateful to the Natural Sciences and Engineer-
ing Research Council of Canada for nancial support
to JLD. We appreciate valuable comments from L.
Fahrig, A. Hegazy, L. Lovett-Doust, H. MacIsaac, G.
Merriam, M. Oldham, W. H. Romme, P. Sale, W.
Reid, and three anonymous reviewers. We acknowl-
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