Habitat Structure, Patchiness, and Avian Communities in North American Steppe

Vegetation: A Multivariate Analysis

Author(s): John T. Rotenberry and John A. Wiens
Source: Ecology, Vol. 61, No. 5 (Oct., 1980), pp. 1228-1250
Published by: Wiley on behalf of the Ecological Society of America
Stable URL: https://www.jstor.org/stable/1936840
Accessed: 26-03-2019 15:11 UTC

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Ecology, 61(5), 1980, pp. 1228-1250
? 1980 by the Ecological Society of America

HABITAT STRUCTURE, PATCHINESS, AND AVIAN

COMMUNITIES IN NORTH AMERICAN STEPPE
VEGETATION: A MULTIVARIATE ANALYSIS'

JOHN T. ROTENBERRY2 AND JOHN A. WIENS

Shrubsteppe Habitat Investigation Team, Department of Biology,
University of New Mexico, Albuquerque, New Mexico 87131 USA

Abstract. We investigated the relationships between vegetational structure, spatial heterogene-

ity, avian community structure, and the ecological responses of the breeding bird populations in the
structurally simple steppe vegetation of North America. We examined the effect of variation in several
measures of both horizontal and vertical patchiness in vegetation distribution on bird distribution and
abundance, then integrated these measures into a smaller subset of variables using Principal Component
Analysis (PCA). We interpreted the new PCA variables as reflective of proximate niche parameters
to which bird species responded.
Out of 550 structure/bird abundance bivariate correlations, 18.5% achieved a significance level of
at least P < .05. Virtually all individual species responses to vegetation habitat measurements were
straightforward: all "typical tallgrass prairie" birds attained highest abundances on plots with highest
percent coverage of grass, "typical shrubsteppe" birds attained their highest abundances on plots
with the highest values of shrub coverage, and so on. In the PCA of vegetational features, the first
axis (which accounted for 41.0o of the total variation) represented variation in horizontal structure
and heterogeneity of vegetation, with high positive factor loadings for 9 of 10 horizontal measures.
The second component (22.4%) represented increasing vertical heterogeneity, with significant cor-
relations for five of five vertical measures. The third component (10.7%) reflected variation in the
density and coverage of low forbs. The segregation of horizontal and vertical measures on orthogonal
axes indicates that variation in horizontal vegetation patchiness is largely independent of variation in
vertical patchiness in these steppe systems.
The abundances of all typically tallgrass prairie bird species were negatively correlated with in-
creasing horizontal heterogeneity and partially correlated with increasing vertical heterogeneity. Sev-
eral shrubsteppe bird species were also positively associated with increasing vertical heterogeneity,
but unlike tallgrass birds their abundances were also positively associated with increasing horizontal
patchiness. Shortgrass prairie birds were generally negatively correlated with vertical patchiness.
Only bird species associated with montane meadows varied significantly with changes in the third
principal component, forb abundance. At the community level, species diversity increased as vertical
heterogeneity increased but appeared to vary independently of horizontal patchiness. Temporal vari-
ation in overall vegetation structure was not closely mirrored by correlated changes in bird community
composition at the shortgrass prairie site for which yearly data were available.
Analysis of species' niche overlaps along the synthetic vegetation axes defined by PCA revealed
a significant lack of niche complementarity; species with high overlap on one axis were quite likely
(P < .001) to have high overlap on a second, independent axis. Analysis of niche metrics across
communities indicated that while average niche breadths of co-occurring species were independent
of a sample's position on any multidimensional axis, average niche overlap increased at the nonmon-
tane grassland sites. Tests of current theories of "diffuse compeition" vs. "niche breadth compen-
sation for low alpha-diversity" were inconclusive. We suggest that the conventional sorts of data
collected to test these hypotheses are inappropriate.
Our results suggest that individual bird species responded independently to the environmental
gradients we derived, and that higher level species interactions were not necessary (and in some cases
statistically improbable) to explain the patterns we observed. These results are consistent with our
contention that even though such biotic interactions may be taking place, they are unlikely to be
detected in our systems because (1) environmental variation obscures relationships between local
populations, (2) populations are not resource limited or equilibrial during large portions of their
history, and (3) individuals are not "fine tuning" their responses to local habitat variation.
Key words: biogeography; birds; community structure; gradient analysis; niche metrics; North
America; principal component analysis; spatial heterogeneity; steppe; vegetation structure.

INTRODUCTION
structuring or physiognomy has repeatedly been
Vegetation gives terrestrial environments their char- shown to be important in determining the distribution
acteristic physical configuration. In turn, this spatial and abundance of birds, acting as a proximate factor
in guiding habitat selection and, presumably, as an
1 Manuscript received 2 August 1979; revised 13 February
ultimate factor by its association with critical re-
1980; accepted 20 February 1980.
sources such as food, nesting sites, or cover from
2 Present address: Department of Biological Sciences,
Bowling Green State University, Bowling Green, Ohio predators (see Hilden 1965 for review). Given such
43403 USA. influences, aspects of vegetational configuration of

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1229

TABLE 1. Features at sample sites shown in Fig. 1. Site numbers refer to Fig. 1.

Number Total
Site breeding of density
Code Grazing Year bird (indi-
Number Name (by plots) treatment sampled species viduals/km2)

Tallgrass prairie
1 Osage OSAG Moderate 1970 4 250
2 Elmdale ELMA Moderate 1967 3 284
ELMB Moderate 1968 4 276
3 Donaldson DNLD Moderate 1968 3 206
4 Fitchburg FTCH Moderate 1966 7 622

Mixed-grass prairie
5 SandhillsSANDA Heavy 1968 3 100
SANDB Ungrazed 1968 5 316
6 Cottonwood COT68 Heavy 1968 4 272
COT70 Heavy 1970 3 153

Shortgrass prairie
7 Pantex PANU Ungrazed 1970 4 212
PANG Heavy 1970 4 267
8 Pawnee PN68A Heavy winter 1968 5 232
PN69A Heavy winter 1969 4 332
PN70A Heavy winter 1970 3 277
PN68B Heavy summer 1968 3 204
PN69B Heavy summer 1969 3 261
PN70B Heavy summer 1970 6 270
9 Laramie LARM Light 1969 2 263

Montane grassland
10 Bridger BRDG Light 1970 4 109
11 Cliff Lake CLFLK Ungrazed 1969 6 551
Shrubsteppe
12 Bison BISN Moderate 1970 3 145
13 ALE ALE1 Ungrazed 1971 3 214
ALE2 Ungrazed 1971 4 291
14 Cabin Lake CABLK Light 1969 4 458
15 Chewaucan CHEW Light 1967 5 140
16 Steens Mountain STEN Light 1967 4 191

habitats are obvious candidates for inclusion as critical
dimensions in analyses of avian niches (e.g., James
1971, Shugart and Patten 1972). The emergence of
multivariate statistical techniques as analytical tools
has fostered an approach to quantification of the mul-
tidimensional Hutchinsonian niche (Green 1971), not
only for birds (James 1971, Anderson and Shugart
1974, Whitmore 1975, Smith 1977), but for a variety
of other organisms as well (e.g., Green 1971 [bivalve
molluscs], Miracle 1974 [plankton], Johnson 1977a, b
[bog plants], Dueser and Shugart 1978 [small
mammals]).
One of the most important components of habitat
structure is spatial heterogeneity or patchiness, which
integrates not only the absolute values of vegetation
characteristics but also their variation in space (Wiens
1976). Other than for vertical foliage height diversity
(e.g., MacArthur and MacArthur 1961, Willson 1974),
whose precise relationship to patch structure is un-
clear, few attempts have been made to relate quanti-
tative measures of patchiness to changes in avian com-
munity composition (but see Wiens 1974b, Roth 1976).
Although some relationship between horizontal and
vertical patchiness is expected, explanations of this
relationship have been imprecise, and no one has yet
successfully merged variation in both planes into one
index of heterogeneity.
Here we take a multivariate approach to describe
habitat niche relations among a small assemblage of
birds in the structurally relatively simple steppe vege-
tation of North America. We describe simple bird/hab-
itat correlations, examine species distributions and
community attributes along gradients in vegetation
structure and heterogeneity in both planes, and esti-
mate niche parameters of each species in a multidi-
mensional environmental space. We then examine
niche relationships among coexisting species and test
predictions of species and community relationships
derived from niche theory.
METHODS
Study areas
Data for this study were collected from a diverse
array of grassland sites scattered throughout middle
and western North America, representing virtually the
entire range of steppe vegetation in this region. Six-
teen sites (Fig. 1) were sampled irregularly during

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1230 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

STEPPE TYPE
Taligrass

Mixed-grass

Shortgrass

~?J Shrubsteppe

FIG. 1. Locations of avian breeding censuses included in this analysis, numerically keyed to Table 1. Steppe types
generalized from Kuchler (1964). Montane grasslands are not shown because of their disjunct distribution.

1966-1971; as some sites had several plots, and some
plots were sampled in subsequent years, a total of 26
samples was available for analysis. Plots included tall-,
mixed-, and shortgrass prairies, montane meadows,
and several Great Basin Artemisia shrubsteppes. A
variety of grazing regimes was superimposed upon this
array of natural vegetation types (Table 1). Sites were
chosen without regard to the number or species of
birds present, but instead reflect our desire to include
a range of structural types sufficient to generate strong
gradients in vegetational heterogeneity.
Data collection
At each site one or more 9.2- to 10.6-ha study plots
were delineated by a staked grid with 61-m grid inter-
vals. All plots were surrounded by a "buffer zone" of
at least 100 m of similar vegetation to reduce edge
effects, and were positioned so as to minimize major
changes in vegetation types within the plot. These
plots formed the basis for both bird population esti-
mation and vegetation sampling.
Bird populations.-Population densities of all bird
species breeding on the study plots were estimated by
mapping individual territories using the "consecutive
flush" procedure (Wiens 1969). These territory maps
not only allowed estimation of population densities but
could also be used to detect which vegetation sampling
points (see below) fell within or outside of a species'
territorial boundaries, permitting a more precise esti-
mation of the species' habitat correlations. These
points were also used to estimate within-plot territorial
overlap, or the average number of species' territories
overlapping at each sample point (Roth 1976). Table
2 lists all species found breeding on the study plots
during the sampling periods. The results of all plot
censuses are available upon request.
Vegetation structure. -Features of vegetation
structure were recorded at sampling units located ran-
domly within each 61 x 61 m block of the sampling
grid. At each sample unit, four sample points were
located at the corners of a square with 2-m diagonals.
Using techniques modified from Wiens (1969), a va-
riety of vegetational attributes was measured at each
point or for each sample unit. The vertical distribution
of vegetation was measured by recording the number
of contacts or "hits" in each decimetre (10-cm) height
interval of a thin (5-mm diameter) rod passed vertically
through the vegetation at each sample point. The type
of vegetation making contact and the depth of litter or
mulch was also noted. The point-centered quarter

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1231

TABLE 2. Breeding birds censused on steppe study sites.

Number Density range

Typical of plots (where present;
Species Code habitat* occupied individuals/kM2)

Mountain Plover (Eupoda montana) MTP SG 1 5

Upland Plover (Bartramia longicauda) UPP TG 4 7-10
Mourning Dove (Zenaida macroura) MDV SG-SS 1 9
Common Nighthawk (Chordelis minor) NHK SG 1 7
Horned Lark (Eremophila alpestris) HLK SG 17 18-197
Rock Wren (Salpinctes obsoletus) RKW SS 1 49
Sage Thrasher (Oreoscoptes montanus) SGT SS 2 15-49
Robin (Turdus migratorius) ROB MM 1 21
Mountain Bluebird (Sialia currucoides) MBB MM 1 16
Loggerhead Shrike (Lanius ludovicianus) LGS SS 1 15
Bobolink (Dolichonyx oryzivorus) BOB TG 1 54
Eastern Meadowlark (Sturnella magna) EML TG 5 74-93
Western Meadowlark (Sturnella neglect) WML MG-SG 16 6-71
Dickcissel (Spiza americana) DCK TG 4 46-91
Savannah Sparrow (Passerculus sandwichensis) SVS TG 2 235-255
Grasshopper Sparrow (Ammodramus savannarum) GRS TG 12 4-185
Henslow's Sparrow (Passerherbulus henslowii) HNS TG 1 25
Lark Bunting (Calamospiza melanocorys) LKB SG 8 2-165
Vesper Sparrow (Pooecetes gramineus) VSP SS-MM 2 7-134
Sage Sparrow (Amphispiza belli) SGS SS 4 57-145
Chipping Sparrow (Spizella passerina) CPS TG-MM 1 19
Brewer's Sparrow (Spizella breweri) BRS SG-SS 6 5-252
White-crowned Sparrow (Zonotrichia leucophrys) WCS MM 1 45
McCown's Longspur (Calcarius mccowni) MCL SG 4 76-150
Chestnut-collared Longspur (Calcarius ornatus) CCL SG 1 62

* TG = tallgrass, MG = mixed-grass, SG = shortgrass, MM

method (Cottam and Curtis 1956) was used to estimate
densities of forbs and woody vegetation using the cen-
ter of each sample unit (intersection of the 2-m diag-
onals) as the point. The height of the forb or shrub
nearest to the point was also measured.
These measurements describe features that fall into
two basic categories: coverage and structural (Table
3). "Coverage" is simply the percent coverage of var-
ious physiognomic classes (grass, forb, shrubs/cacti,
bare ground/rock, and litter), and was obtained from
the frequency of occurrence of these types at all sam-
ple points within a plot. Also included in this category tural heterogeneity. To measure such variation we
are the total density of all forbs and woody vegetation used the coefficients of variation (CV) of total hits
(i.e., shrubs and cacti) and the density of woody plants (CVTOTHIT), of the maximum height interval with
alone, estimated from the point-quarter method.
"Structural" variables have the additional property of
"dimension;" that is to say, variation in a structural
index is generally associated with spatial variation in
either a horizontal or vertical plane. For example, the
average number of hits in the first 10-cm height inter-
val (HITlO; Table 3) or the average litter depth (LIT-
DEP) is associated with vegetational variation in the
horizontal plane, while the average total number of
hits per sample point (TOTHITS) varies more as a
function of the changing vertical distribution of vege-
tation. Other expressions of vertical vegetation struc-
ture include the average maximum decimeter height
interval recording vegetation contacts (MAXHGT),
the average height of the nearest emergent forb or
shrub (TOTHGT), or the average effective height
(EFFHGT-the height to which the surrounding vege-
tation obscures view of a board 3 cm wide). While
these measures are not direct indices of vertical patch-
iness (see below), it is clear that the magnitude of these
means is strongly associated with their variation and,
hence, indirectly reflects spatial variation in vertical
structure.
As the sampling points were arrayed over horizontal
space within a plot or area occupied by a species,
the between-point variation in these vertical struc-
tural variables measures or indexes horizontal struc-
hits (CVMAXHGT), of average forb or shrub height
(CVTOTHGT), and of litter depth (CVLITDEP),
based upon all sample points.
At this point we should note the problem of scale
that arises when we attempt to define "heterogeneity"
or "patchiness" or even vegetation structure using the
above measures. As we have based these measures on
means and variances over all sample points, we have
described heterogeneity on the scale of ==60 m (the
mean distance between sample units). While this scale
is perhaps appropriate for estimating variation on the
order of individual territories, we also desire some
measures of variability approximating the scale of a
localized activity area of an individual. To do this we
follow the approach of Wiens (1974b), which defines
a heterogeneity index based on within-sample-unit dif-

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1232 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

TABLE 3. Description of vegetation structural variables measured on steppe study plots. See text for details of measurement
and calculation.

Range of observed values

Variable Code Min Max

Coverage
1. Percent cover grasses GRASSCVR 0 lOo
2. Percent cover forbs FORBCVR 0 53%
3. Percent cover shrubs/cacti WOODCVR 0 40o
4. Percent cover bare ground/rock BARECVR 0 54%
5. Percent cover litter LITCVR 2 83%
6. Total forb + shrub density TOTDENS 26.9 3431.5/M2
7. Shrub density WOODDENS 0 436.4/M2

Structural-vertical
8. Total number of hits TOTHITS 1.8 9.3 hits
9. Maximum decimetre interval containing hits MAXHGT 0.8 3.6 dm
10. Effective height EFFHGT 0.4 18.0 cm
11. Average height of emergent forbs and shrubs TOTHGT 6.4 40.1 cm
12. Forb + shrub height heterogeneity index HGT-HI 0.16 0.76

Structural-horizontal
13. Total number of hits in first decimetre HIT10 0.7 5.4 hits
14. Litter depth LITDEP 0 3.6 cm
15. Coefficient of variation of total hits CVTOTHIT 0.4 2.1
16. Coefficient of variation of MAXHGT CVMAXHGT 0.3 1.5
17. Coefficient of variation of LITDEP CVLITDEP 0.8 3.7
18. Coefficient of variation of TOTHGT CVTOTHGT 0.4 0.9
19. Coefficient of variation of Max-Min
emergent forb/shrub height CVHGTDIF 0.50 1.35
20. Total hits heterogeneity index HITS-HI 0.79 2.85
21. Litter depth heterogeneity index LIT-HI 1.38 4.80
22. Point-to-plant distance heterogeneity index DIST-HI 0.05 1.07

ferences (a scale of =2 m) averaged over an entire ation of the average Max - Min for emergent forb
plot. Thus, for any attribute we wish to measure at a and shrub heights (CVHGTDIF).
sample point, a heterogeneity index may be defined as
Environmental predictability

H (Max - Min) ? - x (Max - Min)
HI= ~~N N YT
where Max = maximum value of the attribute within
a sample unit, Min = minimum value of that attribute
within the unit, N = the total number of sample
points, and x = the mean value of the attribute within
a sample unit. This within-sample-unit variation in at-
tributes is weighted by dividing by the overall average
of that attribute, on the premise that the biological
importance of a given amount of variation is inversely
related to the magnitude of the overall average (e.g.,
a 2-cm difference in vegetation height is likely more
important in an area with an average vegetation height
of 4 cm than an average height of 20 cm).
The within-sample-unit attributes we have indexed
in this fashion are total hits (HITS-HI), litter depth
(LIT-HI), distance to nearest forb or shrub (DIST-HI),
and height of nearest forb or shrub (HGT-HI); the first
three are horizontal indices, while the fourth is a ver-
tical index. DIST-HI is similar in concept to the spatial
heterogeneity index derived by Roth (1976), while
HITS-HI has been used by Wiens (1974b). As a final
horizontal index, we calculated the coefficient of vari-
Climatic instability has been suggested to limit op-
portunities for niche diversification (Pianka 1974,
MacArthur 1975) and thus is of concern here. We used
time-series autocorrelation (Kendall and Stuart 1967)
to analyze predictability of rainfall regimes. Employ-
ing data from Wiens (1974a), we calculated autocor-
relation of monthly precipitation with lags ranging
from 1-12 mo, with lengths of record ranging from 30-
60 yr. In essence, this estimates the likelihood that
rainfall in any 1 mo will be very similar (or very dis-
similar) to rainfall in any of the 12 previous mo. This
approach thus looks at climatic predictability by ex-
amining the monthly correlations of rainfall amounts
both within seasons and from one year to the next. To
reduce the information from these correlations to a
single index for each site, we calculated their standard
deviation based on the 12 mo. Thus a high standard
deviation indicates substantial departure from zero
autocorrelation and, hence, relatively high predict-
ability; a low standard deviation indicates the oppo-
site. This measure of predictability is independent of
both average annual rainfall and its coefficient of vari-
ation at these sites (r = .20, p > .35; r = .01, p >
.90, respectively).

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1233
Statistical methods
Descriptive statistics for the bird communities sam-
pled on each plot were calculated according to Hill
(1973). Diversity was measured as
N2 = I/ Pt2,
where pi is the proportion of the ith species in the
sample. Richness was expressed as the number of
species observed in the sample. Evenness, E, was
calculated using the ratio proposed by Hill (1973):
E = N21N1,
where
N, = exp(-Ypilnpi).
We have elsewhere (Rotenberry 1978) provided justi-
fication for the use of this series of measures.
While many variations on the general theme of
multivariate manipulation have been used to elaborate
niche structure, most studies rely largely on either dis-
criminant function analysis (DFA) (e.g., Green 1971,
1974, James 1971, Dueser and Shugart 1978) or prin-
cipal components analysis (PCA) (e.g., Miracle 1974,
Johnson 1977a, b), with the technique chosen being
largely determined by the investigator's conception of
what niche axes ought to represent. Those studies di-
rected toward determining the ecological "separa-
tion" of species or the characterization of their eco-
logical differences chose DFA. The rationale for such
a choice has been stated by Green (1974), who, fol-
lowing Levins (1968), avers that niche dimensions are
not the number of biologically relevant factors in the
environment, but the number of factors that can best
be used to separate species. DFA is clearly the opti-
mum technique for identifying variables that contrib-
ute to differences between species. The approach of
searching for niche differences, however, is subject to
conceptual limitations. Traditional niche theory pos-
its, in essence, that species differ in niche features.
Because the power of DFA is in its ability to produce
linear combinations of variables that maximize differ-
ences, it is predisposed to show (and emphasize)
species differences. It thus does not provide an objec-
tive "test" of the theory, and in fact contributes to
the trivialization of niche theory embodied in the state-
ment that different species are, in fact, different, if one
looks hard enough (Wiens 1977a).
PCA, on the other hand, may to a considerable ex-
tent avoid such difficulties by representing what is es-
sentially the approach of gradient analysis (Whittaker
1967, 1975). Sites are directly ordinated along a mul- as
tifactorial gradient defined by PCA, and the abundance
distributions of species plotted and compared. While
such a technique avoids focusing on (i.e., maximizing)
species differences, it nonetheless allows them to be
estimated, based on the reciprocity of their distribu-
tions. While avoiding the pitfalls of DFA, it still meets species x and y in the ith interval (Stander 1
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the two criteria any niche quantification method must
meet (Shugart and Patten 1972): (1) it provides a meth-
od of testing for niche overlaps among species; and
(2) it expresses the size of a species' niche (niche
breadth) in a simple manner but with minimum loss of
information. For these reasons, we employed PCA in
our analyses.
The patterns present in the 26 sites x 22 vegetation
variables matrix were unraveled by means of PCA per-
formed on the 22 x 22 vegetation variable correlation
matrix. This matrix is identical to a variance/covari-
ance matrix of standardized variables (Cooley and
Lohnes 1971). This procedure reduces the number of
environmental variables from the original 22 to a
smaller, more manageable set of orthogonal variables
(components) that still accounts for a large part of the
total variation in the environmental data set. To the
extent that the original variables are highly correlated
with the resulting components (high factor loadings),
these axes may be interpreted in ecological terms, and
the sites (along with individual species' abundances at
those sites) may be ordered along the gradients defined
by these axes.
This use of PCA differs from so-called "indirect gra-
dient analysis" (Nichols 1977) in that we are con-
structing environmental axes (=gradients) directly
from environmental data (e.g., Johnson 1977a, Roten-
berry 1978) rather than inferring environmental axes
indirectly from species distributions. The problems
associated with nonlinear or discontinuous species re-
sponses (Austin 1976) are largely circumvented by the
use of continuously distributed, standardized environ-
mental variables.
Methods similar to those of Johnson (1977b) were
employed to estimate niche breadths and overlaps.
Each principal component axis was divided into five
equally spaced intervals, without regard to the number
of sample plots that fell into each interval. It was
straightforward, then, to estimate the relative abun-
dance of a species across intervals based on its total
abundance in plots within the intervals. A species'
niche breadth was estimated by the diversity of its
distribution (a variance measure) along a resource (=
component) axis, and in our case is calculated accord-
ing to the simple diversity formula previously pre-
sented. Now, however, pi is the proportion of that
species' total abundance that occurs in the ith interval.
As a measure of niche width, this index was originally
proposed by Levins (1968). Niche overlap between
two species was estimated by the similarity of their
distributions along a resource axis, and was calculated
0= E xiyi
VI X,2 >2y,2
where xi and yi are the relative abundance
1234 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

IH-WISI* * * *

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1235
1974, May 1975). This index ranges from zero (no over-
lap) to one (complete overlap), and is symmetrical for
any pair of species.
Relations between two sets of variables were tested
using product-moment correlation coefficients, while
comparisons between means were carried out using
the t' test (Sokal and Rohlf 1969), which requires no
assumption of homogeneity of variance nor equal sam-
ple sizes. The convention adopted throughout the fol-
lowing text is that * implies statistical significance at
the .05 level, ** at the .01 level, and *** at the .001
level. All correlations and multivariate analyses were
performed using the Statistical Analysis System
"SAS76" (Barr et al. 1976). The confounding effects
of covariation between two variables and their inde-
pendent relationships to a third were removed by use
of partial correlations (Steele and Torrie 1960).
RESULTS AND DISCUSSION
Bivariate correlations
The initial step in extracting bird/vegetation rela-
tions is to examine the correlation matrix between
both sets of variables (Table 4). Out of 550 possible
correlations, 18.5% are * or better, 9.6% are ** or
better, and 5.5% are ***. These percentages of signif-
icant values are far higher than those expected by
chance, strongly suggesting that most of the described
relationships are not spurious.
Several patterns begin to emerge at this level. For
example, there was a clear association between the
abundances of Dickcissels, Eastern Meadowlarks, and
Grasshopper Sparrows (typical tallgrass prairie species)
and variables describing tallgrass plots-dense grass
and deep litter (TOTHITS, MAXHGT, LITDEP, LIT-
CVR, EFFHGT, and GRASSCVR). On the other
hand, there were strong negative relationships with
BARECVR and several other variables associated
with increasing horizontal heterogeneity (CVTOT-
HIT, HITS-HI, LIT-HI, and CVHGTDIF). The fourth
typical tallgrass species, Upland Plover, did not dem-
onstrate these relationships as strongly.
Covarying in a different manner are several of the
shrubsteppe species, particularly Brewer's and Sage
Sparrows, Sage Thrashers, and Loggerhead Shrikes.
These species evidenced high positive correlations
with coverage values of woody plants and bare
ground, and with several indices relating to horizontal
heterogeneity, but, conspicuously, not with the den-
sity of woody vegetation (i.e., shrubs). As grass cov-
erage increases, the abundance of these species de-
creases.
Another species showing a definite pattern is the
Horned Lark, with strongly negative relationships to
several vertical indices (MAXHGT, EFFHGT, and
TOTHGT). Other shortgrass prairie species, how-
ever, exhibited less clear patterns; McCown's Long-
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spur responded much less strongly to the same ver-
tical indices and showed a positive relationship with
WOODENS (in this case, Opuntia cactus on several
of the heavily grazed sites on which the longspur was
abundant).
Table 4 indicates that every vegetation variable
measured was significantly associated with the abun-
dance of at least one, and usually four to five, bird
species. The existence of these correlations sug-
gests at the least that these variables are relevant to
the sorts of questions we are asking. GRASSCVR,
BARECVR, and HITS-HI showed perhaps the
strongest patterns, serving to separate the responses
of the tallgrass species (lots of grass, little bare ground,
low horizontal heterogeneity) from those of the shrub-
steppe species (little grass, extensive bare ground,
high horizontal patchiness). Another pattern was
shown by FORBCVR, which was highly correlated
with those species that reached their maximum abun-
dances at the montane sites (Vesper, Chipping, Sa-
vannah, and White-crowned Sparrows, Mountain
Bluebirds, and Robins). These mountain meadows are
often thickly carpeted with wildflowers in the spring,
and while these bird species may be responding to
other features of the environment, their association
with the forbs is nonetheless significant.
Beyond these relations of individual species' re-
sponses to vegetational habitat parameters, the eco-
logical literature suggests that aspects of overall avian
community structuring, such as diversity, should be
closely related to vegetation structure (MacArthur
1958, 1972, MacArthur and MacArthur 1961). Most
causal explanations of this relationship draw attention
to the role of environmental complexity, patchiness,
or heterogeneity in promoting niche diversification,
and thus diversity (Willson 1974, Roth 1976), and in
fact such relationships have been explicitly empha-
sized to explain species numbers in grassland habitats
(Cody 1968). As such heterogeneity is measured in
some of the vegetation variables we considered, we
might expect to be able to define some key habitat
components that enhance diversity in our system. Ex-
amining the straightforward bivariate correlations,
however, adds little to our understanding. Species di-
versity was significantly positively correlated only
with MAXHGT (**), EFFHGT (*), and FORBCVR
(**). This observation is similar to that made by Cody
(1968), who found that he could predict diversity using
average grass height and its variance. Although it has
been suggested that species diversity should decline
as environmental "noise" (i.e., unpredictability) in-
creases (MacArthur 1975), we detected no significant
relationship between diversity and our measure of cli-
matic predictability (r = -.24, P > .25). We also
found no correlations between species richness or
evenness considered separately and any environmen-
tal variables.
1236 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

TABLE 5. Factor loading of vegetation principal components. Only those values that are significantly correlated (P < .05)
with a component are shown; only factors with eigenvalues > 1 are shown.

Factor: I II III IV V
Eigenvalue: 9.02 4.94 2.35 1.40 1.03
Vegetation % a: 41.0 22.4 10.7 6.3 4.7
variable L o% : 41.0 63.4 74.1 80.5 85.2

Coverage
GRASSCVR -0.87
FORBCVR 0.40 0.76
WOODCVR 0.80
BARECVR 0.92
LITCVR -0.84
TOTDENS 0.94
WOODDENS 0.45 0.74
Structural-vertical
TOTHITS 0.56 0.56
MAXHGT 0.77
EFFHGT 0.86
TOTHGT 0.88
HGT-HI 0.70 0.40
Structural-horizontal
HITIO 0.46 0.44 0.64
LITDEP -0.71 0.52
CVTOTHIT 0.83
CVMAXHGT 0.84
CVLITDEP 0.80
CVTOTHGT 0.50
CVHGTDIF 0.68 0.46
HITS-HI 0.93
LIT-HI 0.69 0.47
DIST-HI 0.73

Vegetation factor analysis

Most species, excepting a few that occurred at only
one site, demonstrated some significant population
variability associated with at least one vegetation
character (Table 4). The perceptive field ornithologist
will find none of these correlations counter-intuitive.
However, correlation does not imply causation, and
we should emphasize that we have not demonstrated
habitat "selection" but have merely pointed out hab-
itat "association." A particular complication is that
many of the vegetation variables themselves are in-
tercorrelated; in this data set, almost half (107/231) of
all pairwise comparisons were significantly (* or great-
er) different from zero. Thus, a bird species may vary
significantly in abundance with respect to one variable
when in fact it is selecting or responding to some other
variable that is highly correlated with the first (e.g.,
the FORBCVR/montane meadow example mentioned
above). One of the virtues of multivariate analysis,
such as the PCA we employ here, is that it extracts
independent patterns of covariation from such inter-
correlations.
Derived gradients.-The component patterns re-
sulting from PCA are summarized in Table 5, which
lists all statistically significant factor loadings for each
vegetation variable. No variable reached a high load-
ing beyond the fifth component, and all components
greater than the fifth had eigenvalues <1. In subse-
quent analyses we restrict attention to the first three
components, which combine to account for almost
three-quarters of the variability in the entire vegetation
data set.
Not only do just three components account for the
bulk of the variability in the original data set, but these
components are also readily interpretable, in a some-
what unanticipated fashion. The first component in-
cludes significant positive values for 9 of the 10 hori-
zontal structure measures. As seven of eight horizontal
heterogeneity indices load high on this axis, it seems
clear that this factor represents increasing horizontal
spatial heterogeneity. This horizontal spatial patchi-
ness is associated with increasing coverage of bare
ground and woody vegetation (largely shrubs), in-
creasing shrub density, decreasing coverage of grass
and litter, and decreasing litter depth. The second
axis, on the other hand, clearly represents a vertical
component, with high positive correlations with all
five vertical structure measures. Of the coverage mea-
sures, this vertical component is associated only with
forb coverage. Interestingly, it shows a strong covaria-
tion with DIST-HI, a heterogeneity index that we ini-
tially presumed reflected horizontal variation (see be-
low). The third component reflects variation in the
density and coverage of low forbs (mainly wildflow-

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1237

2.00
ELMA

CHEW

CABLK

z OSAG
z
Lxi 1.00 DNLD

0 ELMS ~~~~~~~~~~~~CLFLK ALE2 STEN

1=FTC ALE
FTCH H~~~ ~~~~ISN
I
0 I~~~~~~~~~~~4
<:V 0 I
(D SANDA

> ~~~~~~~~~~~~SANDS

set8A ( l T5lgrlss
-1.00 COT7OT.~ PN708 LARM A Mixed-gross
PN695 C)C Shortgrass
PN68 P69AShrbstpp

- .0 - II5 .0-05 .010 .020

0

1.00

2.00

-2,00 -1.50 -1.00-00 00o0.50 1.00 1,50 2.00

VEGETATION COMPONENT I

FIG. 2. Site factor scores based on vegetation principal components. See text for a complete interpretation of the axes.
The axes are scaled by the relative contribution of each component in accounting for variation in the total vegetation data
set (Table 5). Plot names are coded as in Table 1.

ers), while the fourth loads high only for the density
of and heterogeneity indices associated with woody
plants. The fifth component is associated only with
variation in the distribution of litter. The increasing
vegetational patchiness observed on the first two prin-
cipal axes is also associated with decreasing climatic
stability (r = -.42* and -.62**, respectively). The
mechanism underlying this observation, that predict-
able steppe habitats are structurally less complex, is
unclear.
Perhaps the most surprising aspect of this multi-
variate vegetational analysis is the distinct separation
of vertical and horizontal components. Although most
studies detailing habitat patchiness/bird species diver-
sity relationships have focused on only one dimension
(e.g., Wiens 1974b for horizontal, Willson 1974 for
vertical), the assumption has always been that the
patch structure in one dimension is generally associ-
ated with that in the other. Our results suggest that
these two dimensions that are perpendicular in phys-
ical space are largely perpendicular (i.e., orthogonal)
in multivariate statistical space, at least for indices
such as those we have calculated. Those few structural
variables that are shared on both axes (TOTHITS,
HITlO, LITDEP) are only weakly reflective of heter-
ogeneity per se. It appears, then, that variation in
vegetational patchiness follows different patterns in
horizontal vs. vertical space, at least in steppe vege-
tation, further implying that the search for a single
index that combines both dimensions (Roth 1976) may
be long and difficult.
The distribution of sample plots along these multi-
variate gradients in habitat structure, heterogeneity,
and forb abundance is illustrated in Fig. 2. These fac-
tor scores are used to quantify a plot's location on
each axis for subsequent correlations with other plot
characters (e.g., species diversity). The tallgrass sites
evidence substantial vertical heterogeneity but rela-
tively little patchiness in a horizontal plane. Shrub-
steppe sites generally show as much vertical hetero-
geneity as the tallgrass ones, but differ markedly in
being much patchier horizontally. Shortgrass sites are
intermediate in horizontal heterogeneity, but, as one
would expect, show very little vertical variability. The
mixed-grass sites are the most varied in their struc-
ture, with Cottonwood and the ungrazed Sandhill plots

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1238 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

~AOSAG,-4

~~4h4~~

B. COT70 D. PN69A

FIG. 3. Photographs of plots that illustrate the extremes represented by the first two axes of the vegetation principal
components (Fig. 2). A. Osage (OSAG)-high vertical, low horizontal heterogeneity; B. Cottonwood (COT70)-low vertical,
low horizontal heterogeneity; C. Cabin Lake (CABLK)-high vertical, high horizontal heterogeneity; D. Pawnee (PN69A)-
low vertical, high horizontal heterogeneity. See Table 1 for plot details.

positioned intermediate to the shortgrass and tallgrass
plots, while the heavily grazed Sandhill plot more
closely resembles the shrubsteppe sites in structure.
The montane meadows are horizontally and vertically
intermediate, but separate from the other sites along
component III, forb coverage and density.
To illustrate the nature of these gradients in the real
world we have included photographs of plots that rep-
resent the extremes (Fig. 3). Osage is a mostly contin-
uous tallgrass sward. Cottonwood is also continuously
covered with grass, but demonstrates little vertical
variability in this coverage. Pawnee also demonstrates
little vertical variability, but is substantially more bro-
ken up horizontally. Cabin Lake has as much or more
vertical relief than Osage, but the resulting patches are
considerably more unevenly distributed horizontally.
Species distributions.-The abundances of several
species of birds are strongly correlated with the PCA
vegetational axes, and those relationships that are sta-
tistically significant are depicted in Fig. 4. This plot
yields fairly distinct clusters of species that respond
numerically to changes in habitat structure in a similar
fashion. All of the species that we have previously
considered "typical" of tallgrass prairies reach their
highest abundances on plots that exhibit lowest hori-
zontal patchiness while exhibiting substantial vertical
development. Shrubsteppe species show a generally
similar response to increasing vertical structure, but
differ antithetically from the former set of species in
their response to horizontal heterogeneity. The short-
grass species do not differ in abundance with respect
to changes in horizontal variability, but all three
shown are significantly negatively correlated with in-
creasing vertical variability. The third vegetation axis
is correlated only with the abundances of two of the
montane meadow species (Robin and Mountain Blue-
bird) and the Vesper Sparrow.
We should note, however, that these correlations
are perhaps less reflective of linear habitat responses
than simply the presence or absence of these species
at one end or the other of the derived gradients. Con-
versely, the absence of a significant correlation does
not necessarily imply the absence of a habitat re-
sponse, for of the 11 species showing no significant
correlation with any component only one, the Western
Meadowlark, occurred on more than one plot. Even

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1239
100
0.50
s | ~~~~~EML
Z | - ~~~~~DCK
Lii
o ~~~~~~~GRS
- 1.00 - 0.75 -0.50
. a 0
Lii
(0D
LuJ
-1.0 0 - _ _ _ _ _ _ _ _ _ _ _ _ _ _
-0.50
0.50
1.00 - _ _ _
-100 0750.0025002
FIG. 4. Correlations between bird species abundances and site factor scores based on vegetation principal components.
The axes are scaled as in Fig. 2. Species names are coded as in Table 2; species that were uncorrelated with any factor (P >
.05) are not shown.
if a one-plot species were strongly affected by some
of the measured habitat variables, the statistical like-
lihood of detecting such a relationship would be small.
Community attributes.-As with the bivariate cor-
relations, we can also examine environment/species
diversity relationships in multivariate vegetation
space. As in so many other studies (e.g., MacArthur
and MacArthur 1961, Karr 1971, Karr and Roth 1971),
we observe species diversity to increase significantly
as vertical complexity increases; the correlation be-
tween diversity and component II is .48*. There is,
however, no detectable relationship to horizontal
patchiness (Component I; r = .11, P > .60), nor to
any other multidimensional vegetation pattern. We
have predicted elsewhere (Rotenberry 1978) that sig-
nificant changes in diversity along a gradient in hori-
zontal spatial heterogeneity should reflect changes in
evenness rather than richness. However, not only did
we detect no change in overall diversity along the hor-
izontal component, but neither of the components of
diversity was correlated with that component (r =
-.13, .02; all P > .50). Willson (1974) predicted that
diversity changes along a gradient in vertical com-
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SGS
SGT
-0.25 0 0.250.500.751.00LGS
BRS
VSP
UTaligrass
0 Shortgrass
* Shrubsteppe
0.50 ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~montane
MBB,
ROB LKB MCL
5 0.75 1.00
VEGETATION COMPONENT I
plexity should be associated with species richness; the
diversity association we noted above with respect to
increasing vertical heterogeneity was not, however,
parallelled by either component (r = .17, -.13; P >
.40). Our data do not constitute a fair test of either
prediction, however, because of the substantial differ-
ence in the scale of measurement; both Willson's and
Rotenberry's gradients included the full range between
steppe and mature forest vegetation, while the data
here encompass only a very small portion of that
range.
Roth (1976) has provided a theoretical framework
in which to look at patchiness/diversity relations that
is more appropriate to the scale of measurement used
here. Roth proposes two alternative explanations for
complexity/diversity correlations: (1) horizontal over-
lap between species does not increase with increasing
species richness, implying that the number of percep-
tible (to birds) patch types increases in the habitat,
and that utilization of them is coarse grained; or (2)
overlap does increase with increasing numbers of
species present, suggesting that increasing heteroge-
neity does not measure the addition of coarse-grained
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1240 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5
patches, and that birds stratify vertically and/or use
other resources (e.g., food) in a coarse-grained way.
Implicit in these alternatives, of course, is the as-
sumption that some resource is in short supply and
that ecological partitioning is necessary to permit
species' coexistence.
Our data certainly lead us to reject alternative (1):
the number of species overlapping at a point is clearly
a function of the number of species (r = .61***) and/
or the diversity of species (r = .49**). Additional
species are accomodated in the community by increas-
ing overlap, not increasing patch segregation. This is
further buttressed by the observation that overlap is
independent of increasing horizontal heterogeneity
(Component I; r = -.28, P > .20). This leads us to
alternative (2), which suggests separation along
another ecological dimension. We have elsewhere sug-
gested that food resources are rarely limiting in grass-
land or shrubsteppe habitats (Wiens 1977b; Rotenber-
ry 1980), and have demonstrated substantial levels
of dietary overlap in the very species considered here
(Wiens and Rotenberry 1979, Rotenberry 1980).
Although we have not explicitly considered other re-
source relationships, it is difficult to imagine what oth-
er resources might be potentially limiting in this sys-
tem.
It has been suggested that where territorial or hor-
izontal overlap is great, ecological separation may be
accomplished by vertical segregation (MacArthur
1964, Cody 1968). While this appears a useful axis for
resource division in a forest or brushland, we question
its feasibility in vegetation that rarely averages over
1 m tall. The activity budget information that we have
gathered (J. T. Rotenberry and J. A. Wiens, personal
observation for species in shrubsteppe, short-, and
mixed-grass prairies) reveals no stratal specialists and,
given the list of species with which we are dealing
(Table 2), there are no intuitively obvious candidates.
Nevertheless, while diversity is correlated with in-
creasing vertical complexity (see above), the number
of species overlapping at a point is essentially random
with respect to this component (r = -.22, P > .90).
But if the number of overlapping species is divided by
diversity (Roth 1976), this new index (representing the
percent of the entire community that overlaps at a
point) varies in a significantly negative fashion with
increasing complexity (r = -.41*), as Roth also ob-
served.
What is clear from the foregoing is that there is sub-
stantial inconsistency in our results when they are
couched in Roth's construct. We observe a corollary
result (decreasing percent overlap with increasing
complexity) without confirming a primary prediction
(independence of overlap and richness). The problem
appears to lie more with the nature of the indices rath-
er than the soundness of the model and its predictions.
We did not use precisely the same index as Roth, but
only one analogous to it, and our index was shown to
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vary more as a vertical than a horizontal measure
(Table 5). We are led to suggest, however, that both
indices in fact measure vertical complexity rather than
horizontal patchiness as Roth and we initially thought.
This likely results from the facts that these indices are
generated from point-to-plant measurements, and that
the plants measured (forbs, shrubs, and trees) contrib-
ute more to vertical complexity than to horizontal
patchiness. This implies that the significant diversity/
complexity correlations observed merely continue the
trend initiated by the MacArthurs (1961).
We are left to conclude that, given the largely or-
thogonal nature of structure and heterogeneity in the
two dimensions, a consistent body of theory relating
changes in bird species diversity to horizontal spatial
heterogeneity is still incomplete. Such incompleteness
may stem from (1) our inability to measure horizontal
patch structure correctly, and on the proper spatial
scale; (2) our inability to assess vertical segregation
correctly; and/or (3) the fact that species in this system
may not be resource limited, and hence spatial overlap
and diversity become random variables with respect
to patch structure.
Niche relationships
Niche breadth.-The multivariate orderings of
vegetational features documented above also permit
an examination of the niche patterns of the bird
species in this vegetation space. The distribution on
the first two vegetation components of each species
occurring on more than one site is plotted in Fig. 5.
The components have been divided into five equally
spaced intervals (cf. Johnson 1977b). The number for
each species in the figure is a measure of the species'
"niche breadth" along that environmental axis, and
is essentially the diversity of its distribution among the
intervals. Species found at only one plot, and hence
in only one interval, would have a niche breadth of
1.00, while species found equally distributed among
all intervals would have the maximum possible
breadth, 5.00. The location of each species along the
component reflects the correlations outlined in the
previous section. The third component is not shown,
but most species were not widely distributed along it,
with niche breadths ranging only from 1.38 to 2.65 for
those species shown in Fig. 5.
The lack of correlation of the abundant Western
Meadowlark with any component is associated with
their wide-ranging, rather general response to the en-
vironmental parameters we have measured; meadow-
larks had the broadest niche of any species on all three
components. Species such as Grasshopper Sparrows
and Horned Larks showed a pattern of relatively wide
niches on one axis but relatively narrow ones on the
opposite (the axis of "specialization"). Other species,
such as McCown's Longspurs, were not widely dis-
tributed with respect to either axis and, as will be
October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1241

0.8- Dickcissel .-.. Eastern Grasshopper Upland Plover Horned Lark Western
Meadowlark Sparrow Meadowlark

W044422.77 227 /70 293 390

o 062
z

> 0 8 Lark Bsnting Brewer's Sparrsw McCown's Longspsr Vesper Sparrow Savannah Sparrow Sage Sparrow

F-J0

04 2.28 2. 178 222 2.00 200

1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5

VEGETATION COMPONENT I

0.8 Dickcissel Eastern Grasshopper Upland Plover Horned Lark Western

Meadowlark Sparrow Meadowlark
0.8

S0.00
0.4- 1.55 2.19 4.03 2.66 1.87 3.99

< 0.2-

1.0
W 1.0 Lark Bunting Brewer's Sparrow McCown a Vesper Sparrow Savannah Sparrow Sage Sparrow
0.8 Longspur

W 08 --
1.90 2.44 1.00 1.92 2.00 ] 2.00

0.2- . 1 ...L.. . . 1 S ,

1 2 3 4 5 1 2 3 4 5 1 -2 3 4 5 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5

VEGETATION COMPONENT 11

FIG. 5. The relative abundance of bird species in each of five equally spaced intervals along the first two vegetation
principal components. The number in each graph refers to the niche width of that species along the axes. See text for details.
Only species that were observed on more than one plot are shown.

shown, had very low overall niche breadth even While uncommon species tended to have small niche
though they were abundant when present. breadths as measured here, the opposite was not al-
Overall niche breadth can be taken to indicate a ways apparent. Within groups of habitat-typical birds
species' performance on several niche dimensions no real trends were apparent. Each group had several
combined. Because the axes described here are by species with minimal breadths of 1.00, several average
definition orthogonal and hence independent, overall
niche width is simply the product of a species' niche
widths on each single axis (May 1975, Pianka 1975).
TABLE 6. Overall niche breadths for those species occurring
These overall widths are presented in Table 6 for those on more than one plot, based on the first three vegetation
species occurring on more than one plot. The maxi- principal components. See text for method of calculation.
mum possible breadth is 5 x 5 x 5 = 125 equally oc-
cupied intervals. Species that were sampled on only Overall
Species niche breadth
one plot have an overall width of 1.00.
Quite clearly the Western Meadowlark was the most Upland Plover 6.74
Horned Lark 12.88
generalized of all the species considered.This is not,
Eastern Meadowlark 4.54
however, a consequence of its "dominance" (sensu Western Meadowlark 41.24
McNaughton and Wolf 1970), for it was neither the Dickcissel 4.86
most ubiquitous species in these samples (the Horned Savannah Sparrow 8.00
Grasshopper Sparrow 15.19
Lark was) nor the most numerically abundant, either
Lark Bunting 8.63
where it occurred (the Savannah Sparrow was) or over Vesper Sparrow 11.76
all plots combined (the Horned Lark was). It did, how- Sage Sparrow 5.52
Brewer's Sparrow 14.58
ever, account for the greatest proportion of bird bio-
McCown's Longspur 3.17
mass on over half the plots on which it was present.

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1242 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5
members with breadths of -5-6, and a more general
member with a width of 12-15. We should emphasize
that these derived gradients appear to be fairly realistic
with respect to species' distributions, and hence are
not unreasonable "niche" axes. Although many of
these species occur at geographically widely separated
sites, most show a continuous distribution along these
gradients. To some extent this is an artifact of dividing
the components into equally spaced intervals, but
even within intervals occurrences tend to be contig-
uous. While the criticism might be made that these
intervals are unweighted with respect to actual utili-
zations by individual species (e.g., Colwell and Fu-
tuyma 1971), other studies have suggested no a priori
reasons to assume Gaussian response curves along all
environmental gradients (see especially Terborgh
1971). Likewise, no species is given a wider niche be-
cause we inadvertently sampled many vegetationally
similar plots on which it was abundant; such plots
would tend to cluster within the same interval.
Niche overlap.-Niche overlap is a measure of the
degree to which two species co-occur in the intervals
defined along the components, and is calculated ac-
cording to the similarity formula presented earlier. An
intuitive grasp of the relationships defined thereby can
be gained from comparison of species' distributions in
Fig. 5. For example, Eastern Meadowlarks and Dick-
cissels were almost identically distributed along the
first component, and their calculated overlap (0.98)
reflected this. Sage Sparrows, however, were distrib-
uted exclusively of those two species along the first
axis, and hence had zero overlap with both. Most
species pairs lie somewhere between these two ex-
tremes. Similar relationships between species can be
seen on the second axis as well.
Overlaps calculated in this manner are of course
confounded by the spatial co-occurrence of species in
plots; such aspects of local and geographic distribu-
tions may be at least partially unrelated to the rela-
tionships of species on the vegetational gradients we
have derived here. For example, for all species that
occurred on more than one plot, species-pair overlaps
on the first niche axis (component) are significantly
correlated with overlap based on spatial co-occurrence
alone (r = .71***). There still exists a substantial
amount of variation in "niche overlap" that is inde-
pendent of "space overlap," however, as the coeffi-
cient of determination (r2) is only .50 (that is, half of
the variation in niche overlap along the first axis is
independent of spatial co-occurrence). Where appro-
priate in the following analyses, the residual effects of
this relationship with spatial co-occurrence will be re-
moved by the use of partial correlations.
Of perhaps greater interest is not merely the overlap
between two species on one niche dimension but their
relationships when several axes are considered simul-
taneously. Conventional niche theory predicts that
species that overlap extensively along one axis should
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diverge strongly along another in order to permit co-
existence (May 1975, Pianka 1975). As these niche
dimensions are truly independent, overall multidimen-
sional overlap is calculated as the product of the sev-
eral separate one-dimensional overlaps (May 1975).
However, although the two axes are by definition in-
dependent from one another, species-pair overlaps on
the first axis are highly correlated with overlaps on the
second when the effects of spatial co-occurrence are
partialled out (rpar, = .59***). Apparently, then, these
species are in fact not independently distributed from
one another in the different niche dimensions. The
mechanisms) that would produce this nonrandomness
is not detectable with these data; however, we shall
offer some speculations concerning its nature in our
concluding remarks.
The niche overlaps of all species that occurred at
more than one site along the first three vegetation
components are graphically summarized in Fig. 6. The
dendrogram was formed by simple weighted pair-
group cluster analysis (Sokal and Sneath 1963) of the
overlap matrix based on the first three axes. Three
distinct groups were distinguished at the .50 overlap
level: birds with affinities for tallgrass, for shortgrass,
and for shrubsteppe habitats. These groups are not
precisely congruent with those we have used in earlier
treatments. For example, Savannah and Vesper Spar-
rows demonstrate niche responses more like tallgrass
species, and Western Meadowlarks were more closely
allied with shortgrass species. However, if we choose
to define a similarity level for group formation that
splits off those three species (i.e., a group-defining
level of .60), then Sage Sparrows, an obvious shrub-
steppe bird, must be segregated from the shrubsteppe
group. Inclusion of all species in construction of the
dendrogram serves mainly to increase the within-clus-
ter heterogeneity without altering any of the basic re-
lationships we have already observed. It should be
pointed out, too, that this dendrogram does represent
multidimensional overlap largely independent of spa-
tial co-occurrence, for the correlation between spatial
overlap and a matrix of overlaps derived from the den-
drogram is only r = .04 (P > .70).
Mean habitat vectors
Another way to synthesize bird/habitat relationships
in a form that can be viewed simultaneously for all
species is Principal Components Analysis of the mean
habitat vector for each of the species (Anderson and
Shugart 1975). Each vector consists of the average
value for each environmental parameter for each bird
species, estimated only from those sample points lying
within a species' territorial boundaries (i.e., areas ac-
tually occupied by individuals within plots). In gen-
eral, the factor loadings and major component axes
resulting from this PCA of mean habitat variables
closely conform to those of the overall site habitat
variables and, in fact, closely duplicate the informa-
October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1243
"OVERLAP"
0 0.50
FIG. 6. Dendrogram of the niche overlap relationships among all species that were observed on more than one plot. Niche
overlap is measured by the co-occurrence of two species in the intervals defined along the first three vegetation PCA axes
(Fig. 5 and text).
tion in Table 5. The first component derived from the
mean habitat vector (o- = 41.6%o) is very similar to the
first component in Table 5; the correlation between
factor loadings resulting from the two analyses is r =
.87***. Likewise, the second (a = 24.1%) and third
(a = 10.6%) mean habitat vector components are very
similar to the second and third site vegetation com-
ponents (r = .87*** and .82***, respectively). The
fourth (a = 8.0%) and fifth (a = 4.8%) components of
the second analysis (the only remaining ones with ei-
genvalues >1) become increasingly less similar (r =
.40* and .32, respectively). The great similarity be-
tween the results of these two analyses strongly sug-
gests that the two data sets being analyzed are in fact
very similar. We infer from this that there is very little
within-plot habitat selection in these species, which
may be a consequence of our attempt to minimize
overall within-plot heterogeneity when initially se-
lecting plot locations.
Species are shown in the mean habitat space (three
dimensions, 76.3% of the total variance) in Fig. 7.
Comparison with habitat/abundance relationships
(Fig. 4) shows little change for those species depicted
in both, although the inclusion of previously uncor-
related species in Fig. 7 reduces the clustering appar-
ent in Fig. 4. We observe no particular dispersion of
species with rather similar ecologies as did Anderson
and Shugart (1974); most species that eat the same
things (Wiens and Rotenberry 1979, Rotenberry 1980),
for example, tend to be close in habitat utilization as
well. It is, however, difficult to separate the cause-
effect relationships involved here; it is unclear wheth-
er these birds eat the same things because they live in
the same places, or live in the same places because of
the availability of certain items. If in fact structural
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1.00
Dickcissel
Eastern Meadowlark
Grasshopper Sparrow
Upland Plover
Vesper Sparrow
Savannah Sparrow
Western Meadowlark
Horned Lark
Lark Bunting
McCown's Longspur
Brewer's Sparrow
Sage Thrasher
Sage Sparrow
habitat selection is the more important variable driving
where a species occurs (as has been stressed through-
out these and other analyses), then it seems likely that
the first alternative proposed is the more logical one.
Temporal variation
Throughout, we have treated spatial heterogeneity
as temporally invariant, changing only from plot to
plot. As the two Pawnee plots were sampled for three
consecutive years, we can use them to examine the
change in multivariate space relations through time.
Fig. 8A, which is a magnification of a small portion of
Fig. 2, shows the degree to which temporal variability
in vegetation is expressed (although this variation is
necessarily less than that exhibited between sites; cf.
Fig. 2). On Pawnee B most of the temporal variation
represented a steady progression toward less horizon-
tal patchiness. Pawnee A, on the other hand, fluc-
tuated in both dimensions, although most of this vari-
ation was likewise concentrated horizontally. While
undoubtedly some part of the difference between the
two Pawnee plots represents the difference in grazing
regimes (winter vs. summer), both plots had similar
stocking rates and had undergone continuous grazing
treatment for many years prior to our sampling (D.
Jameson, personal communication).
Did bird populations mirror these temporal shifts in
vegetation structure? The changes in Pawnee plot bird
composition is best shown, as for vegetational
changes, in multidimensional space. Fig. 8B represents
the location of 3 yr of Pawnee censuses along principal
component axes defined by the bird species data sets
for both plots. Rather than representing "indirect gra-
dients" per se, these axes merely define patterns of
covariation in species' abundances (Nichols 1977).
1244 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5

LOS
2.00

1.00

LU EML WCS Tollgrass

- 1.0-10 -05 .HN0 0 Short1r0ss
BOB ~~~~~~~~~~~~~~~~~~~~~~~~~~~~0Shrubsteppe
C.)
o A ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~montane
ov RKW

T GRS SGT

BBS

LUV CPS

SOP

-1.00

MOO

CCL LKB
MBB MTP MC

0 1 1 t00

NHK

(~~~~~~~~~~O 1.00 { ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1.00

0.00 200

-1I.50 -1.00 -0.00 0 00;0 1060 1.50 0.00 2.500

VEGETATION COMPONENTI

FIG. 7. Bird species factor scores based on principal component analysis of mean habitat vectors. See text for a comp
interpretation of the axes. The axes are scaled by the relative distribution of each factor in accounting for variation i
total mean habitat data set. Species names are coded as in Table 2.

The first axis (52.4% of the total variation) represents
a dine from high abundance of McCown's Longspurs
and Homed Larks (positive end) to high abundance of
Lark Buntings and Brewer's Sparrows. The second
axis (21.6%) has high positive loadings for Homed
Larks and a high negative loading for Western Mea-
dowlarks, while the third (14.8%) weakly contrasts
longspurs (low) with meadowlarks (high). Their com-
bined variance is 88.8% of the total.
Statistical comparison of the location of the plots in
multivariate vegetation space and multivariate "bird
space" reveals little in the way of associated trends
between the two. Of the nine correlations possible be-
tween plot/year factor scores based on vegetation axes
I, II, and III, and bird axes I, II, and III, only one
(bird I/vegetation III) was significant (r = -.87*); as
forb coverage and density increased, buntings and
Brewer's Sparrows increased while larks and long-
spurs decreased. The average for the nine was F = .49
(average P > .35). In general, then, changes in vege-
tation structure were not closely matched by changes
in the avian community. Assuming that the bird PCA
is an appropriate description of the multispecies com-
munity, we suspect that these observations stem from
a lack of fine tuning of birds to their "niche gestalt"
(sensu James 1971). Moreover, to the extent that any
of these species display site tenacity, fine-scale envi-
ronmental tracking may be even more blurred. Birds
may return to the same territory in subsequent years
despite changes in habitat features, especially if their

FIG. 8. Temporal (yearly) variation on two shortgrass prairie plots (Pawnee). A. Variation in vegetation structure based
on factor scores of vegetation principal components. The axes are scaled as in Fig. 2, but here only that portion of the
multivariate space including these shortgrass sites is shown, magnified =4x. B. Variation in bird communities based on
factor scores of avian community principal components. The axes are scaled by the relative contribution of each factor in
accounting for variation in the total Pawnee bird census data set. See text for details and interpretation of the axes.

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October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1245

A
-0.50

LLi -0.75

PN68A

iLPN70B P PN68B

-0.25 0 0.25 0.50 0.75

VEGETATION COMPONENT I

B 9
1.00i
A~~~~~~ PN69A~~~~~~~~~~N8
PN77 A

A, '\-05

0 2568 PN02569B .
BIRDAIO COMPONENT I

-I5

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1246 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5
previous breeding attempt was successful (Hild6n
1965). What we see in any one season will be less a
function of a close degree of habitat selection as the
environment varies but more an expression of survi-
vorship and reproductive output of the previous year.
All of this is to suggest that while many species may
show the appropriate responses on the regional level
(within-site variation at Pawnee is much less than be-
tween-site variation over this entire study), there still
exists fuzziness at the local level (within-site variation
remains). This may be expected to obscure predicted
relationships between coexisting populations and sug-
gests a lack of environmental fine tuning.
Niche relationships between communities
The determination of the niche parameters of width
and overlap in multivariate space enables us to ex-
amine how these parameters vary with respect to each
other, the position of communities along the derived
vegetation gradients, and the number of species in a
community. In particular, we can test predictions
about niche relations that have been made from con-
siderations of resource partitioning, diffuse competi-
tion, and environmental predictability. The methods
we use are similar to those of Johnson (1977a, b).
We should note first that there is a strong negative
relationship between the average three-dimensional
vegetational niche breadth and the average niche over-
lap of all species coexisting at any given site (r =
-.49**). In other words, the pattern at a site is for co-
occurrence of relatively generalist species that other-
wise overlap little in habitat utilization outside the site,
or, conversely, for co-occurrence of specialist species
that also share the same specialities (i.e., have high
niche overlap). Both imply conditions of nonlimiting
resources with subsequent opportunistic exploitation
of those resources by coexisting species, conditions
we have elsewhere argued as being normal for grass-
land bird populations (Wiens 1974a, Rotenberry 1980).
This inverse relationship is, however, at least partly
confounded by inadequate sampling for some species;
that is, a species may be characterized as narrow
niched not because its spectrum of vegetation struc-
ture utilization is narrow, but because only one or a
few of our sample plots fell within that species' oth-
erwise large biogeographical range. Such seems likely
to be the case for Robins, Mountain Bluebirds, Com-
mon Nighthawks, and White-crowned Sparrows, for
example. On the other hand, the relatively low ob-
served niche breadths of Mountain Plovers, Hen-
slow's Sparrows, and Bobolinks probably do reflect
a rather narrow range of habitat specificity. The ex-
ample of the Bobolink points out, too, that relative
habitat specialization (certain types of tallgrass prai-
ries [Tester and Marshall 1961]) need not result in geo-
graphic restriction, especially if such habitat is wide-
spread.
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Neither average niche width nor average niche over-
lap of coexisting species is correlated with a site's po-
sition along the axis of horizontal heterogeneity (r =
.18 and -.20, both P > .30), and the partial correla-
tions are even worse (r = .10 and -.13, both P >
.50). (In the following examinations of relations among
niche metrics and various site attributes, we will at-
tempt to reduce the effects of the inverse niche
breadth/overlap relation by use of partial correlation.)
Likewise, we observe no absolute correlation between
niche metrics and either of the other two PC vegeta-
tion axes (r = -.33, -.13, -.09, -.37 for breadth and
overlap on axes II and III, respectively). However,
partial correlations reveal two significant patterns.
First, statistically controlling for overlap yields a sig-
nificant partial correlation between average niche
breadth and vertical heterogeneity (r = - .46*);
species coexisting at vertically more complex sites
tend to have narrower overall habitat niches. The sec-
ond and perhaps more interesting pattern is that when
variations in niche breadth are partialled out, niche
overlap among species is significantly negatively cor-
related with position on axis III (r = -.48**); sites
that are toward the negative end of this axis have
species with high niche overlap. Reference to Fig. 2
indicates that the majority of the true grassland (i.e.,
nonmontane) sites have negative values for Factor III.
Pianka (1974) has pointed out that extensive niche
overlap is likely associated with reduced competition;
if resources are not in short supply, organisms can
share them without detriment to one another. We have
suggested elsewhere that resources are rarely in short
supply in typical grassland and shrubsteppe habitats
(Wiens 1974a, 1977b, Rotenberry 1980) and this result
is consistent with that suggestion.
Cody (1975) has proposed that resource-limited as-
semblages of birds where habitat overlaps are high
should evidence separation in another dimension; if
this alternative axis is food, he predicts a correlation
between increasing habitat overlap and increasing sep-
aration in body sizes. Taking masses from Wiens
(1971) and R. E. Ricklefs (personal communication),
we calculated average ratios between adjacent pairs
of birds on the body size spectrum in each plot. How-
ever, we found no correlation between this ratio and
those species' average habitat overlap (r = -. 12, P >
.50). We have shown elsewhere (Wiens and Roten-
berry 1979, Rotenberry 1980) that body size (or bill
length) is a poor predictor of either the size or taxa of
food taken by members of this same set of grassland
birds; these results, when combined with the above
indication of resource nonlimitation, suggest that the
basic premise of Cody's speculation, that body size
separation should index trophic niche divergence, is
suspect.
Elements of the "niche overlap hypotheses" of
Pianka (1974) have been associated with the concept
October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1247
of "diffuse competition" (MacArthur 1972, Pianka
1974), ahnd lead us to examine the relationship between
clusions, deriving from his model the prediction that
these niche metrics and species diversity. Pianka pre-
dicts that under conditions of diffuse competition,
niche overlap should vary inversely with respect to
diversity, while niche breadth should remain relatively
unchanged. Under conditions of classical resource
partitioning, however, one expects to find what Cody
(1975) describes as niche breadth compensation for
low alpha-diversity: under conditions of low diversity,
niche breadths should be broad, narrowing as diversity
increases. Niche overlap, on the other hand, is ex-
pected to remain unchanged.
Using the same methods as Pianka and Cody we
find that our data may be used to support (or refute)
either hypothesis; the partial correlations indicate that
both niche breadth (r = -.57**) and niche overlap
(r = -.43*) decrease as species diversity increases.
However, these correlations do not indicate that in-
dividual species have "adjusted" their niche metrics
in the face of changing competitive pressures. We sug-
gest that it is much more likely that data such as ours
are largely inappropriate for a proper test of these hy-
potheses because species' niche widths and overlaps
are estimated from geographical data, rather than at
the population (i.e., within-site) level, where compet-
itive interactions presumably take place. Our gener-
alizations from the higher to the lower level may be
confounded by a variety of interpopulation interac-
tions (e.g., niche compression) that are simply unde-
tectable between sites. We feel that these hypotheses
have been inadequately tested by Cody (1975) and
Pianka (1975), for the same reason. The most straight-
forward interpretations of such results are that (1) the
species present at sites with high diversity tend to have
narrow niches, while those at sites with low diversity
tend to have broad niches; or (2) average overlap is
less among species at high-diversity sites than at low-
diversity ones. We must stress, however, the statis-
tical independence of the observations that results
from the use of partial correlations. These results do
not imply that species at high-diversity sites have nar-
row, nonoverlapping niches while those at low-diver-
sity sites are broadly overlapping generalists. They do
indicate, however, that within sets of sites of species
with similar breadths, overlap decreases as diversity
increases; likewise, within sets of sites of species with
similar overlaps, average breadth decreases as diver-
sity increases.
Finally, we can examine variation in the niche met-
rics of coexisting species at sites that vary with respect
to climatic predictability. For example, Levins (1968)
has predicted that wide-niched species are better
adapted to uncertain environments; likewise Slobod-
kin and Sanders (1969) suggested that environmental
unpredictability selects for organisms with wide tol-
erances. Approaching the problem from another di-
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rection, Leigh (1975) came to essentially the same con-
increased specialization rather than increased niche
overlap will characterize species in more constant en-
vironments. May and MacArthur (1972) also predict
that niche overlap should not increase as predictability
increases.
Correlations from our habitat data show that species
existing at more climatically predictable sites have
greater average overlap among themselves than those
at less predictable sites (r = .41*), while their niche
breadths seem independent (r = .15, P > .45). Partial
correlations indicate that both overlap (r = .56**) and
width (r = .44*) increase significantly as climatic pre-
dictability increases. If we assume that productivity
or habitat quality and environmental stability are as-
sociated on the scale that we measured it, then these
results are in accordance with what we have elsewhere
described as an opportunistic fine-grained response to
abundant nonlimiting resources (Wiens and Rotenber-
ry 1979). As the abundance of resources increases,
species responding in a fine-grained manner will dem-
onstrate relatively broad niches. As most of the
species present will be capitalizing upon the same set
of resources, their mutual overlaps will be high also.
We suggest that most of the results presented above
are consistent with this interpretation of these species'
responses throughout the steppe habitat.
CONCLUDING REMARKS
Our results have clearly shown correlations between
the abundances of many species and physiognomic
features of their habitat, both for univariate (Table 4)
and multivariate (Figs. 4, 5) characters. Although we
fully recognize that correlation does not imply caus-
ation (habitat selection, in this case), we infer that
birds are responding either to the variables we have
measured or at least to unmeasured features that are
strongly associated with those variables. Given that
most considerations of the effects of spatial hetero-
geneity have been focused on community structure
(MacArthur 1964, Willson 1974 for vertical heteroge-
neity; Wiens 1974b, Roth 1976 for horizontal), we find
the fact that individual species apparently respond to
gradients in patchiness and structure (Figs. 4, 5) of
interest. This means that we are not constrained to
think of variation in community structure along such
gradients as reflecting higher level interactions among
species that co-occur at a site (although we certainly
cannot preclude their existence), but are free to ex-
plore alternative hypotheses that may account for the
species assemblages we observed.
The synthesis of avian responses to both horizontal
and vertical heterogeneity in vegetation structure has
led to observations that were not immediately pre-
dictable from the results of previous efforts that were
mainly concerned with the horizontal or vertical di-
1248 JOHN T. ROTENBERRY AND JOHN A WIENS Ecology, Vol. 61, No. 5
mension alone. For example, it was generally antici-
pated that variation in patchiness in one dimension
was correlated with variation in the other; the results
of the PCA clearly indicate that such is not the case
for the indices we have used. Conventional theory also
predicted that, given two independent niche dimen-
sions, high overlap between two species along one axis
would be "compensated" by reduced overlap along
the second; the observed high positive partial corre-
lation between all pairwise overlaps on vegetation
axes I and II is strong evidence that such is not the
case in our system of birds, at least for these dimen-
sions.
Indeed, the common thread that emerges from vir-
tually all of our analyses is the apparent lack of close
interactions or biotic coupling between coexisting
species. We have argued elsewhere that assemblages
of grassland and shrubsteppe birds are not likely to be
equilibrial, largely because climatic variability serves
to prevent species abundances from rising to the point
where resources become limiting or, alternatively,
provides frequent conditions of resource superabun-
dance (Wiens 1974a, 1977a, b; Wiens and Rotenberry
1979; Rotenberry 1980). In neither case are competi-
tive relationships expected to be important on a year-
to-year basis.
While our data suggest that intracommunity struc-
ture is not a function of close biological relationships
among coexisting species, we nevertheless do observe
clusters of species and communities along our derived
gradients that appear different (that is, we have "typ-
ical" tallgrass species, "typical" shrubsteppe species,
etc.). We suspect that most of these species tend to
follow more or less independent distributions with re-
spect to one another, but are drawn together by their
common response to similar features of the habitat.
The broad-scale patterns we see are modified from the
pure Gleasonian individualistic distributions by the
very real presence of environmental discontinuities
(Terborgh 1971, Whittaker 1975), which are evidenced
by the fact that site clusterings are apparent (Fig. 2)
even though PCA attempts to define statistically con-
tinuous gradients in multivariate space.
Much of the evidence we offer in support of our
position that these species are largely decoupled from
direct interactions with one another is negative; that
is, many of the relationships predicted from theoretical
considerations of community structure based on com-
petition and subsequent resource partitioning are not
apparent in our collection of data. We suspect the fail-
ure of these predictions is a result of a breakdown in
the assumptions of the models applied, assumptions
that are too often accepted without close examination
(Wiens 1977a). In particular, we suggest that the im-
plicit conditions of resource limitation are seldom met
and that, in general, we are dealing with nonequilibrial
populations that exist in habitats that are temporally
variant.
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More important, however, are those statistically sig-
nificant relationships that we observe that are counter
to conventional predictions. The example of correla-
tions between overlaps on orthogonal axes is partic-
ularly compelling. Neither did we observe any partic-
ular overdispersion of ecologically similar species in
the multidimensional space defined by the mean hab-
itat vectors; species with similar ecologies tended to
be found in similar habitats. This observation, based
on within-plot data, was further supported by the be-
tween-plot results, which indicated that species at typ-
ical grassland sites tended to have high multidimen-
sional niche overlap.
There is yet a third class of results we observed,
those that were statistically significant but gave sup-
port either to a competition-oriented or to a biologi-
cally decoupled explanation of the data. For example,
our examination of Roth's model of spatial overlap/
species diversity/habitat heterogeneity falls into this
category. We observed that while the percent spatial
overlap among species decreased with increasing
vegetational complexity, the number of species over-
lapping at a point appeared to be a direct function of
the number of species present; the first result implies
some sort of biotic compensation (Roth 1976), the lat-
ter some large element of biotic independence. Also
falling into this third class of results are the observa-
tions from the partial correlations of niche widths and
overlaps with species diversity.
The prevalent approach to understanding commu-
nity patterns applies data taken from broad geograph-
ical areas to test hypotheses that are founded upon
interactions between individuals and populations: for
example, in a recent book devoted to the explication
of community patterns using arguments based on in-
terspecific competition (Cody and Diamond 1975),
most of the evidence offered to support those argu-
ments came from comparative biogeographic studies.
A potential difficulty in this approach is that obser-
vations gathered over broad geographic regions are
used to draw inferences about mechanisms, such as
competition, producing large-scale patterns. But the
processes ultimately responsible for such distribution-
al phenomena (predation, birth and death schedules,
physiological limitations, etc., as well as competition)
go on among individuals and populations at a local
level, and the most appropriate observations with
which to examine community hypotheses would thus
seem more likely to come from examining population
dynamics at the within-site level than from broad geo-
graphic comparisons. In our systems, however, the
detection of such processes at a local scale is unlikely
because (1) populations are unlikely to be resource
limited or equilibrial during large portions of their his-
tory, (2) individual populations may not be fine tuning
their responses to subtle habitat variations, and (3)
differences in patterns of environmental variation be-
tween different areas may obscure relationships be-
October 1980 HABITAT STRUCTURE AND BIRD COMMUNITIES 1249
tween different local populations. We thus find a per-
plexing mixture of agreement with prevailing theory,
disagreement with theory, and ambiguity. We suggest
that this dilemma is more apparent than real, and that
our perplexing mixture is likely a consequence of the
fact that no single mechanism is responsible for com-
munity patterns. As a consequence, we strongly urge
that future biogeographical analyses pay particular at-
tention to alternative mechanisms that may be equally
consistent with the patterns detected.
ACKNOWLEDGMENTS
Roy Mason and Dave Niess were involved in early de-
velopments of these analyses, and Glenn Ford, Peter Grant,
Fran James, and an anonymous reviewer drew our attention
to several inaccuracies in an earlier draft of the manuscript.
The field data were gathered during studies supported by
National Science Foundation grants GB-6606 to J. A. Wiens
and GB-7824, GB-13096, GB-31862X, GB-31862, and
DEB73-02027 A03 to the Grassland Biome, United States
International Biological Program. Data analysis was sup-
ported by the computer centers of Oregon State University
and the University of New Mexico, and the final stages of
this research were supported by National Science Founda-
tion grant BMS 75-11898.
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Austin, M. P. 1976. On non-linear species response models
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Barr, A. J., J. H. Goodnight, J. P. Sall, and J. T. Helwig.
1976. A user's guide to SAS76. SAS Institute, Raleigh,
North Carolina, USA.
Cody, M. L. 1968. On the methods of resource division in
grassland bird communities. American Naturalist 102:107-
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. 1975. Towards a theory of continental species di-
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