Ecological Correlates of Insular Rodent Diversity

Author(s): Raymond D. Dueser and William C. Brown

Source: Ecology, Vol. 61, No. 1 (Feb., 1980), pp. 50-56
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1937154
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Ecology, 61(1), 1980, pp. 50-56
? 1980 by the Ecological Society of America

ECOLOGICAL CORRELATES OF INSULAR

RODENT DIVERSITY'

RAYMOND D. DUESER
Department of Environmental Sciences

AND

WILLIAM C. BROWN
Department of Biology, University of Virginia,
Charlottesville, Virginia 22903 USA

Abstract. Rodent populations were censused on nine Virginia barrier islands during the summers
of 1975 and 1977. The five species observed include Microtus pennsylvanicus, Mus musculus, Ory-
zomys palustris, Rattus norvegicus, and Sciurus carolinensis. The number of rodent species ranged
from zero on a small, low-lying, sandy island to five on a large, elevated, forested island. Rodent
species richness (diversity) increased directly with island area, elevation, vegetation height, and
habitat complexity (i.e., the number of distinct woody plant associations). Because of strict zonation
of plant associations, increased habitat complexity represents increased availability of exploitable
patches of habitat. As expected from studies of mainland rodent communities, this patchiness appears
to promote increased species diversity. Each species exhibits an "exploitation specialty" involving
little spatial overlap with other species. Although no direct tests have been conducted, we speculate
that the rodent communities of these islands are at equilibrium. Low rates of colonization and high
rates of extinction interact to produce low species diversity and a steep species-area curve.
Key words: barrier island; colonization; extinction; island biogeography; Microtus pennsylvan-
icus; Mus musculus; Oryzomys palustris; Rattus norvegicus; rodents; Sciurus carolinensis.

INTRODUCTION The number of vascular plant species is a good pre-

Preston (1962a, b) proposed that the number of
species (S) on an island is related to its area (A) by
the expression S = bAz, where b and z are the inter-
cept and slope, respectively, of the species-area curve.
Preston (1962a) also calculated a theoretical "expect-
ed" z value of ~0.27, based on the assumptions that
species abundances follow a lognormal distribution
and that the total density of individuals is approxi-
mately constant from island to island. Because of its
obvious generality, this species-area model occupies
a central position in equilibrium theories of island bio-
geography (MacArthur and Wilson 1963, 1967). Sev-
eral investigators, however, have examined the influ-
ence of factors other than area on insular species
diversity, both to determine the effects of particular
proximate factors and to explain deviations of ob-
served z values from the expected value of 0.27. While
these studies have been limited in the types of equi-
librium predictions they could test (Simberloff 1974,
1976a), they have identified variables which facilitate
regional predictions of S for particular taxonomic
groups (e.g., Diamond and Mayr 1976).
For insular vertebrates, the apparent importance of
island area generally diminishes as more explicit mea-
sures of environmental complexity are considered.
I Manuscript received 20 April 1978; revised March 1979;
accepted 7 May 1979.
dictor of breeding bird diversity on the Galapagos Is-
lands (Harris 1973), on the high, rocky California
Channel Islands (Power 1972, 1976), and on the low,
sandy Hawaiian Islands (Amerson 1975). Plant diver-
sity (but not shrub volume diversity) also accounts for
much of the variation in lizard species diversity on
islands in the Gulf of California (Case 1975). Habitat
diversity, measured as the number of gross habitat
types and the number of coniferous tree species, is a
good predictor of breeding bird diversity on montane
islands in the Great Basin of the western United
States. The present study examines the relationship
between several physical and habitat variables and the
number of small mammal species on the Virginia bar-
rier islands. Our results demonstrate the influence of
habitat complexity on small mammal diversity (rich-
ness) and suggest that these communities exhibit a
form of species equilibrium.
STUDY AREA
The Virginia barrier islands extend 87 km along the
seaward margin of the Delmarva Peninsula. South-
ward from Wallops Island through Fisherman Island
there are 12 barrier islands, seven bayside islands, and
extensive marshes and marsh islands. The islands are
separated from the mainland and from one another by
broad channels and bays, and are accessible only by
boat. Although several of the islands were sparsely
inhabited prior to 1940 (Graham 1976), the United
February 1980 INSULAR RODENT DIVERSITY
States Coast Guard Station on Parramore Island is
now the only occupied dwelling. Except for intermit-
tent livestock grazing on several islands, the islands
of this complex effectively have been abandoned for
the past 40 yr. We live-trapped small mammals on
eight barrier and four bayside islands. Since storm
conditions cut short our trapping on one barrier and
two bayside islands, we report data for only nine is-
lands. These nine islands encompass the range of area,
elevation, and vegetation complexity found in the
complex south of Wallops Island.
Vegetational differences among these nine islands
are most pronounced in the variety and spatial extent
of woody plant associations. Much of Parramore Is-
land supports mature Pinus taeda forest, mixed pine-
deciduous forest, and tall (>3 m) shrub thickets dom-
inated by Juniperus virginiana, Myrica cerifera, and
several other species. Smith Island is similar to Par-
ramore in vegetational complexity, but lacks extensive
pine forest. Revel Island supports a zone of mixed
forest and low (<3 m) shrub thickets dominated by
Juniperus and Myrica. Hog Island supports sparse
Ammophila breviligulata grassland, dense Spartina
patens grassland, and extensive zones of tall Myrica
thicket. Wreck and Myrtle Islands are predominantly
mixed grassland, but each supports a zone of tall My-
rica thicket. Cobb Island supports sparse Ammophila
grassland and dense Spartina grassland. Little Cobb
supports sparse Ammophila grassland. Mink Island is
mostly marsh, with a patch of mixed grassland and
low shrub thicket dominated by Baccharis halmifolia
and Iva frutescens. All of the islands have extensive
fringing marsh dominated by Spartina patens and S.
alterniflora. McCaffrey (1975) provides a detailed
vegetation map and description for each island except
Skidmore and Raccoon.
SAMPLING
As on other islands along the mid-Atlantic coast,
the vegetation on the Virginia barrier islands exhibits
conspicuous zonation. A transect from the beach to
the bayside marsh intersected representative areas of
most plant associations and topographic features on
each island. Transect locations were selected by
ground reconnaissance following inspection of in-
frared aerial photography and the McCaffrey (1975)
vegetation maps. On Parramore, 200 trap stations
were located along one straight line; on each of the
other islands two parallel lines were established 60 m
apart. One Sherman trap (7 x 9 x 23 cm) was placed
every 15 m along each line, and one Tomahawk wire
trap (15 x 15 x 40 cm) was placed at every 10th trap
station. All traps were baited with rolled oats and
cracked corn, and checked both morning and evening
for four consecutive days. Trapped animals were ear-
tagged for permanent identification and released at the
point of capture. In 5,119 trap nights we compiled 735
captures of 545 individuals of Oryzomys palustris (rice
51
rat), Mus musculus (house mouse), Microtus penn-
sylvanicus (meadow vole), and Rattus norvegicus
(Norway rat). Sciurus carolinensis (gray squirrel) was
observed on two islands but was never captured
(Dueser et al. 1979).
Variation in the rates at which species are captured
is a potential source of conservative bias in any esti-
mate of mammal species composition based on short-
term sampling (Smith et al. 1975). Two pieces of
information suggest that our estimates of species num-
ber (S) are accurate. First, there is a positive but non-
significant correlation between the number of species
captured and both the number of trap stations on an
island (r = .53, P > .20) and the number of trap
nights (r = .64, P > .10). Second, recent observa-
tions from an extensively sampled island indicate that
our procedures for locating transects and for trapping
reveal species composition efficiently. Assateague Is-
land is a large (7029 ha), heterogeneous island 35 km
north of Parramore Island. Paradiso and Handley
(1965) reported seven species of small mammals on
Assateague, including the four species captured on the
Virginia barrier islands plus Cryptotis parva (least
shrew), Peromyscus leucopus (white-footed mouse),
and Zapus hudsonius (meadow jumping mouse). In
August 1977, R. D. Dueser (personal observation)
captured each of these species except Cryptotis during
only two nights of trapping on a 90-station transect
selected by the procedures used here. Only species
listed by Paradiso and Handley (1965) were captured
in over 10000 trap nights on 30 additional transects.
THE VARIABLES
The dependent variable S (species diversity) is the
number of terrestrial rodent species observed on an
island. Eight independent variables are considered
(Table 1). Island area (A), including contiguous marsh,
was determined by planimetry of United States Geo-
logical Survey (USGS) 1:2400 topographic maps (1968
series). Island length (L) is the size of the "target"
provided by the island for animals moving seaward
from the mainland. Isolation from the mainland (I) and
maximum elevation (E) also were determined from
USGS maps. Since these nine islands are arranged
linearly from north to south but are not contiguous, it
seems appropriate to measure isolation as distance
from the mainland rather than as distance between
islands. Maximum vegetation height (H) was deter-
mined directly by selective sampling, using infrared
aerial photographs and vegetation maps to locate ma-
jor vegetational zones. The numbers of vascular plant
associations (V) and woody plant associations (W)
were taken from the vegetation maps prepared by
McCaffrey (1975). Trapping was conducted in all of
the major and most of the minor plant associations on
each island. The number of emergent soil types (T)
was determined from the soil survey maps prepared
52 RAYMOND D. DUESER AND WILLIAM C. BROWN Ecology, Vol. 61, No. 1

TABLE 1. Ecological characteristics of nine islands in the Virginia barrier island complex.

Vascular Woody
Vegeta- plant plant
Isola- Eleva- tion associ- associ-
Species Area Length tion tion height ations ations Soil types
(#) (ha) (km) (km) (m) (m) (#) (#) (#)
Island S A L I E H V W T
Parramore 5 2197 12.8 7.7 9.1 16 15 6 9
Revel 3 508 5.2 6.3 3.0 12 11 3 4
Myrtle 3 407 3.5 9.6 3.7 3 5 1 6
Smith 3 512 7.3 4.7 2.1 9 14 4 6
Hog 2 1177 11.4 11.0 3.0 5 12 3 8
Cobb 1 532 8.3 12.1 2.1 2 9 1 5
Wreck 1 370 5.0 9.8 2.7 4 6 2 5
Mink 1 305 2.4 6.9 1.2 2 4 1 2
Little Cobb 0 29 1.4 9.9 0.9 1 2 0 2

by the United States Soil Conservation Service (G.J.
Hennessey, editor, 1976).
RESULTS
The diversity (number) of small mammal species
ranged from zero on Little Cobb to five on Parramore
(Table 2). All of these species were rodents. The num-
ber of islands on which a particular species was ob-
served ranged from two for Rattus and Sciurus to
eight for Oryzomys. The presence of Mus and Rattus
may be due to past human activity on some of the
islands. The frequency of co-occurrence for each pair
of species is too small to be tested statistically. Never-
theless, there is strong agreement between observed
and expected frequencies of co-occurrence for most
pairs, suggesting that each species is distributed
among islands independently of the other species (Ta-
ble 3). Furthermore, since each species on the average
occurred on only 3.8 of the nine islands, the islands
appear to be effectively isolated and/or significantly
different from one another ecologically.
Correlation coefficients measuring the relationship
between rodent species diversity and each of the eight
independent variables were calculated for both un-
transformed and log,,,-transformed data (Table 4). Cor-
relations for the untransformed data are comparable
to those for the transformed data. The number of ro-
dent species (S) is positively and significantly corre-
lated with area (A), elevation (E), vegetation height
(H), vascular plant associations (V), woody plant as-
sociations (W), and soil types (T) (P S .05). Several
of these independent variables, in turn, are signifi-
cantly correlated. In particular, E, H, V, W, and T all
increase with increasing A. The variables S, A, E, H,
V, W, and T are negatively correlated with isolation
(I) from the mainland. There is a trend for reduced
variety of plant associations and fewer mammal
species with increased distance from the mainland.
Using S or log S as the dependent variable, two
forms of simple regression were computed: linear
using untransformed data, and curvilinear using log10-
transformed data (Table 5). The coefficient of deter-
mination is higher for the curvilinear species-area re-
lation (r2 = 69W) than for the linear (58%). The slope
(z + s,) of the logarithmic species-area curve is 0.38 ?
0.09. There also are significant curvilinear regressions
with L, E, H, V, W, and T, and linear regressions with
E, H, V, W, and T. The coefficients of determination
are somewhat higher for the curvilinear regressions.
Nevertheless, from inspection of the residual variance
in each regression, the data appear to satisfy the as-
sumptions of the linear model (Draper and Smith
1966), indicating that S increases linearly with these
variables. Among these variables, elevation (E, r2 =

TABLE 2. Small mammal species on nine islands in the Virginia barrier island complex.

Microtus
Oryzomys Mus pennsyl- Rattus Sciurus Total Trap
Island palustris musculus vanicus norvegicus carolinensis species nights
Parramore X X X X X 5 2115
Revel X ... X X ... 3 330
Myrtle X X X ... ... 3 290
Smith X X ... ... X 3 495
Hog X X ... ... ... 2 652
Cobb X ... ... ... ... 1 367
Wreck X ... ... ... ... 1 385
Mink X ... ... ... ... 1 160
Little Cobb ... ... ... ... ... 0 315
Islands occupied 8 4 3 2 2 ... 5109
February 1980 INSULAR RODENT DIVERSITY 53

TABLE 3. Frequencies of co-occurrence of particular species TABLE 5. Summaryof simple linear and curvilinearregres-
pairs. Values in parentheses along the diagonal are the sion analyses. r2 = coefficient of determination.P = sta-
number of islands on which a species occurred. Lower tistical significanceof regressioncoefficient(t test).
triangular matrix gives observed frequencies, upper trian-
gular matrix the expected frequencies, under the assump-
tion that the distributions of the species among islands are Linear Curvilinear
independently determined. (S) (log S)
Independent
variable r2(%) P r2(%) P
Oryzo-
mys Mus Microtus Rattus Sciurus
Area A 58 69 .01
Length L 37 48 .05
Oryzomys (8) 3.5 2.6 1.8 1.8 Isolation I 21 20
Mus 4 (4) 1.3 0.9 0.9 Elevation E 72 .01 73 .01
Microtus 3 2 (3) 0.7 0.7 Vegetationheight H 77 .01 79 .01
Rattus 2 1 2 (2) 0.4 Plant associations V 58 .05 65 .01
Sciurus 2 2 1 1 (2) Woody associations W 73 .01 73 .01
Soil types T 55 .05 57 .05

72%), woody plant associations (W, r2 = 73%), and

vegetation height (H, r2 = 77%) are particularly im-
Johnson 1975), rodents (present study) and, perhaps
portant in explaining variation in the number of rodent
less clearly, lizards (Case 1975).
species found on an island.
We infer that increased habitat complexity facili-
DISCUSSION tates increased rodent species diversity through in-
creased resource availability and/or increased habitat
Habitat complexity
stability. Other things being equal, greater resource
Each of the independent variables A, E, H, V, W, availability and greater habitat stability should act to
and T explains at least 50% of the observed variation decrease rates of extinction and perhaps even to in-
in the number of rodent species, even with a small crease rates of colonization, thus increasing species
sample of islands. The number of rodent species oc- diversity. These same effects might be expected from
curring on an island increases with the area and ele- an increase in the area of an island. Using experimen-
vation of the island, and with the height of the vege- tal manipulations of very small (<0.2 ha) mangrove
tation, the number of plant associations, and the islands, Simberloff (1976b) demonstrated that area has
number of soil types. Correlations between the inde- an effect on the number of arthropod species which is
pendent variables confound interpretation of the in- independent of the effect of "habitat diversity." Un-
dividual effects of these variables on the dependent fortunately, it is impossible to specify the individual
variable (S). This is particularly true for those vari- effects of area and habitat complexity with the present
ables for which the coefficients of determination fall data, because of correlations between A, W, V, and
within a narrow range (say ?5%) of variation. None- H. Therefore, the plausibility of our inference that
theless, it is clear that the measures of habitat com- habitat complexity exerts a primary influence on these
plexity W (r2 = 73%) and H (r2 = 77%) have at least rodent communities depends in large part on corrob-
as much predictive power as the physical factors E oration by studies of small mammal communities else-
(r2 = 72%) and A (r2 = 58%). The supposition that where.
increased habitat complexity promotes increased In this respect, the habitat variables which appear
species richness is now supported by studies of insular to be particularly important (i.e., W and H) are those
birds (Power 1972, 1976, Harris 1973, Amerson 1975, which logically should influence the diversity of in-
sular small mammals. Vegetation structure is an im-
TABLE 4. Correlation (r) between number of rodent species portant determinant of the inter- and intrahabitat dis-
(S) and eight ecological variables (defined in Table 1 and tributions of most small mammal species. Ecologically
the text). Lower triangular matrix gives untransformed similar species generally exhibit pronounced habitat
data, upper triangular matrix the log-transformed data.
Italicized coefficients are statistically significant at P S .05 differences in sympatry, with the local distribution of
(two-sided test, df = 7). each species centered in a different habitat (Grant
1972, 1978). Similarly, the distributions and abun-
S A L I E H V W T dances of these species within a habitat are influenced
S ... .83 .69 - .44 .85 .89 .81 .85 .77
by factors such as foliage profile, shrub volume, and
A .76 ... .91 -.12 .83 .76 .89 .86 .82 percent ground cover (e.g., Rosenzweig 1973,
L .61 .88 ... - .01 .75 .72 .94 .83 .89 M'Closkey 1975, 1976, Dueser and Shugart 1978).
I -.46 -.04 .10 ... -.02 -.55 -.32 -.47 .06
E .85
Both habitat selection and interspecific competition
.91 .69 -.10 ... .78 .71 .77 .86
H .87 .74 .61 - .56 .78 ... .86 .96 .68 for space have been observed to influence patterns of
V .76 .74 .87 -.34 .60 .81 ... .92 .81 habitat exploitation by insular rodents (Crowell and
W .86 .84 .79 -.47 .78 .93 .91 ... .76 Pimm 1976).
T .74 .84 .89 .09 .77 .58 .78 .77 ...
Against this background, it is not surprising that
54 RAYMOND D. DUESER AND WILLIAM C. BROWN
measures of vegetation complexity appear to have a
strong influence on insular rodent diversity. Variables
such as W and H may reflect opportunities for effec-
tive habitat segregation between species. Each plant
association comprises a discrete patch of habitat
which differs from the others in species composition,
dominant life-form, and topographic position (Mc-
Caffrey 1975). An increase in the number of plant as-
sociations, therefore, represents an increase in the
number and variety of patches of habitat. Paralleling
this increase in habitat complexity is a similar trend
in rodent diversity, from zero to one species on grass-
land islands, to two or three species on islands having
extensive shrub thickets or forest, up to five species
on the most heavily forested island. Although the
species overlap somewhat in their occupancy of plant
associations, there still is habitat segregation between
species on each island (Brown 1977).
The variables W, V, and H probably describe dif-
ferent aspects of habitat complexity. W (r2 = 73%)
includes relatively stable patches of habitat with
shrubs or trees, while V (r2= 58%) includes these
patches plus the salt marshes and low-lying grasslands
which are subject to periodic innundation. Thus W
may be more important than V in accounting for vari-
ation in rodent diversity because woody plant asso-
ciations contribute heavily to both long-term resource
partitioning between species and high carrying capac-
ities. Both resource partitioning and high carrying ca-
pacities should positively influence rodent diversity.
We assume that H (rW= 77%), like W, is a correlate
of habitat stability in this physically rigorous environ-
ment. In addition, increasing vegetation height may
promote greater species richness through increased
vegetation stratification. Such stratification must be
critical for the addition of an arboreal species such as
Sciurus and may be important even to other species.
Species equilibrium
Only experimental studies (e.g., Crowell 1973, Sim-
berloff 1976a, b) or long-term observations (e.g., Dia-
mond and May 1977) can provide unambiguous tests
of equilibrium theories of island biogeography. Our
assessment of the status of the small mammal fauna
of the Virginia barrier islands is therefore speculative.
The least-squares estimate of the slope of the species-
area curve for this fauna (z = 0.38) is much greater
than the "equilibrium" value of 0.27 hypothesized by
Preston (1962a). In fact, even allowing for statistical
variation, our observed z is slightly above the range
of values (0.18-0.33) generally observed for vertebrate
taxa occupying "isolated universes" (see May 1975,
and Diamond and May 1976, for review). Neverthe-
less, given the restrictive assumptions on which it is
based, the equilibrium value of 0.27 provides a ques-
tionable basis for judging the status of a biota. We
interpret the relatively steep species-area curve ob-
served here as evidence that factors other than area
Ecology, Vol. 61, No. 1
promote increased species diversity, rather than as
prima facie evidence of nonequilibrium.
MacArthur and Wilson (1967) hypothesized a bal-
ance between recurrent rates of colonization and ex-
tinction for equilibrium biotas. Three independent
pieces of information suggest that colonization rates
must be relatively low for rodents on the Virginia bar-
rier islands. First, species known to be abundant on
the adjacent mainland (e.g., Peromyscus leucopus and
Blarina brevicauda) are absent from the islands. Sec-
ond, Microtus, Mus, and Rattus are absent from sev-
eral islands on which habitat conditions appear to be
favorable. Third, three islands which were flooded at
least as recently as March 1962 (the Ash Wednesday
Storm) now support conspicuously depauperate ro-
dent faunas; Little Cobb has zero species, Cobb and
Mink one species each. Any resident populations
probably went extinct while these low-lying islands
were flooded for most of two tide cycles during this
storm. Only Oryzomys palustris, a semiaquatic rat,
now occupies these islands; other species apparently
have not colonized successfully during the past 17 yr.
Conversely, three independent pieces of informa-
tion suggest that extinction rates must be relatively
high for rodents on these islands. First, Mus persists
on only four of the islands and Rattus on only two, in
spite of the almost inevitable introduction of these
species to most of the islands in the past. Second, as
mentioned above, species diversity is conspicuously
low on those islands which are particularly susceptible
to flooding. Catastrophic extinction probably is not an
unusual event on the low-lying grassland and marsh
islands. Third, Bailey (1946) mentioned reports of
Microtus pennsylvanicus on Smith Island and Pero-
myscus leucopus on Hog Island. We trapped Microtus
on the mainland and on three islands, but not on
Smith; Peromyscus was the most abundant species
captured on the mainland, but was never trapped on
an island. Assuming that our sampling was sufficient
to detect Microtus on Smith and Peromyscus on Hog,
these species are no longer present on these islands.
We conclude that these rodent communities are at
equilibrium. Low rates of colonization and high rates
of extinction interact to produce low species diversity
and a steep species-area curve. Although we cannot
discount the independent importance of island size,
our results suggest that these communities approach
saturation with respect to habitat complexity. Because
of the naturally low diversity of the mainland (source)
fauna, this equilibrium is likely to be static rather than
dynamic, with species turnover rates very close to
zero. Crowell (1973) reached similar conclusions about
the status of the rodent communities on small islands
in the Gulf of Maine. He found that islands a few hect-
ares in size support equilibrium communities com-
posed of a single species, and the species turnover rate
is zero. Communities on large land bridge islands con-
tain three species, while those on large, remote islands
February 1980 INSULAR RODENT DIVERSITY
contain no more than two. Furthermore, Crowell
found that the distributions of the three most ubiqui-
tous species are consistent with expectations based on
the propagule survivorship model of MacArthur and
Wilson (1967). The insular distribution of each species
is largely explained by its particular combination of
demographic attributes and colonization ability.
Brown (1971) reached very different conclusions for
mammalian faunas on montane islands in the Great
Basin. He concluded that the number of species in a
montane fauna does not represent an equilibrium be-
tween the rates of colonization and extinction. These
boreal mountaintops were connected to species-rich
faunal sources until the late Pleistocene, permitting
colonization by a diverse group of species. Current
colonization rates are effectively zero because
"bridges" of montane habitat no longer exist. Extinc-
tions have not yet reduced the number of species to
the point where colonizations and extinctions are in
balance, especially on the larger islands. The rates of
colonization and extinction interact to produce an ex-
ceptionally steep species-area curve (z = 0.43). As
further evidence of the nonequilibrium status of these
faunas, Brown (1971) found no significant correlation
between the number of species and any of several
measures of isolation. These faunas thus depart in sev-
eral ways from expectation based on the MacArthur
and Wilson (1967) equilibrium model of island bio-
geography.
More recently, Brown (1978) found no significant
correlation between the number of mammal species
and the measure of habitat diversity devised by John-
son (1975) in his analysis of the avian faunas of the
same montane islands. Brown attributed the low cor-
relation to the dependence of this index of habitat di-
versity on the presence or absence of particular tree
species and habitat types. The index may be an in-
appropriate measure of habitat complexity for mam-
mals. From this result, Brown concluded that area,
elevation, and isolation are the variables best suited
for constructing and testing biogeographic theory.
Even though their effects on species diversity may be
indirect and imprecise, these variables facilitate gen-
eralizations which transcend the peculiarities of par-
ticular taxonomic groups. Brown did grant, however,
that analysis of specific factors which influence the
insular diversity of particular taxonomic groups can
provide information on the mechanisms and effects of
the underlying processes of colonization and extinc-
tion.
MacArthur (1965) suggested that insular mammal
communities often may depart from equilibrium, par-
ticularly under conditions of difficult colonization. The
montane faunas studied by Brown (1971, 1978) appear
to fit this expectation because of their relatively ex-
treme isolation. The rodent faunas of the coastal is-
lands of Maine and the Virginia barrier islands appear
not to fit this expectation, presumably because they
55
are less isolated. Based on our results, we suggest that
any insular mammal community which does exhibit
long-term equilibrium will be saturated with species
sensu MacArthur and Wilson (1967) relative to habitat
complexity and/or stability. Crowell and Pimm (1976)
observed a strict competitive hierarchy for the three
common species on the islands of Maine. Further-
more, they determined that the ability of a species to
occupy its preferred habitat in the presence of other
species corresponded to its position in this hierarchy.
Although we lack such direct evidence of competition,
we hypothesize that competition is similarly important
in sorting species among exploitable patches of habitat
on the Virginia barrier islands, thus creating a strong
positive relationship between rodent diversity and
habitat complexity.
ACKNOWLEDGMENTS
R. M. Dueser, G. S. Hogue, and J. H. Porterassisted with
the fieldwork.This study was possible only because of the
vegetation maps and field assistance provided by C. Mc-
Caffrey. The Nature Conservancyprovidedaccess to the is-
lands of the Virginia Coast Reserve and Conservancy per-
sonnel assisted with field logistics. M. Castagnagenerously
made availablethe facilities of the EasternShore Laboratory
of the VirginiaInstiute of Marine Science. R. K. Rose, D.
Simberloff, and an anonymous reviewer commented con-
structively on the manuscript.This research was supported
by a Universityof VirginiaInstitutionalGrantto R. D. Dues-
er.
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