1 Habitat structure:
The evolution and
diversification of a complex
tOpIC
E. D. McCoyandS. S. Bell
1.1 'HABITAT STRUCTURE' IN ECOLOGY
Habitat structure, by definition, is a component of every ecological study.
This book deals with a particular type of structure, that provided by the
arrangement of objects in space. Even restricted in this way, habitat
structure conjures up a multitude of images in the minds of ecologists, from
concrete topographic features to near-abstractions like 'patches', 'mosaics',
and 'gradients'. The variety of types of physical habitat structure has, in
turn, spawned a wealth of narrowly defined terms meant to convey subtle
aspects of the relationship between organism and structure. While these
terms may do exactly what was intended of them, we suggest that the gain in
precision is offset by a loss in generality. The various subdisciplines of
ecology adopt terminologies, and experimental techniques related to them,
largely for the cognoscente. 'Profile of vegetational density' and 'canopy
surface structure', for instance, may end up having explicit meaning for a
particular ecologist, whereas 'substrate heterogeneity' and 'enemy free
space' may not. Yet, all reside under the broad umbrella of habitat
structure. The outcome of this scenario is that from study to study, one has a
difficult time determining, for example, if 'spatial heterogeneity',
'architectural diversity', and 'habitat physiognomy' are equivalent concepts;
or if 'height density profile', 'foliage height diversity', and 'horizontal
layering' are interchangeable terms; or what unifying features exist among
measurements such as 'soil particle size', 'crown architecture', and 'colony
geometry'. We think this problem gets in the way of comparisons among
studies, and thereby may mask interesting trends; and that it reduces the
recognized importance of physical habitat structure as an ecological agent.
Table 1.1 illustrates the lack of generality which is responsible for our
concern, by showing some studies of vertebrates interacting with a single
S. S. Bell et al. (eds.), Habitat Structure
© Chapman and Hall 1991
4 Habitat structure
Table 1.1 Ways in which 'habitat structure' was measured in a dozen studies of interactions
between vertebrates and vegetation
Study Measurement
Cody and Walter (1976) vegetation height, vegetation density at
different heights
Vince et al. (1976) distance between plants, plant density
Cody (1978) vegetation density at different heights
Dueser and Shugart (1979) vertical woody and herbaceous foliage
density, shrub density, understorey tree
dispersion, overstorey evergreenness,
stump sizes and density
Nilsson (1979) eight vegetation variables
Ewald et al. (1980) percentage cover sufficiently tall to
obstruct vision
Sabo (1980) eighteen vegetation variables
Wittenberger (1980) vertical vegetation density, vegetation
biomass
Wiens and Rotenberry (1981) percentage cover, horizontal and vertical
vegetation density, vegetation spatial
variation, litter depth
Hansson (1982) slash cover compared to brush cover
Karr and Freemark (1983) plant density, percentage foliage cover at
different tree/shrub and understorey
heights, litter depth
Erdelen (1984) vegetation presence/absence at different
heights
type of structure, vegetation, that use the same term, 'habitat (foliage,
vegetational) structure', for structural measurements. Even within this very
narrow grouping of studies, little agreement seems to exist as to precisely
what 'habitat structure' means. The same conclusion also appears to follow
for other terms. For example, Moran (1980) uses the terms 'architecture'
and 'structural complexity' for measurements of plant height and cladode
