11 Habitat structure and

community dynamics in
marine benthic systems
K. P. Sebens

Animals and plants of the marine benthos depend on their substrata for
attachment, on surfaces or within sediments, and for access to a source of
prey and nutrients. In the case of sessile species, the two-dimensional
surface is a primary resource for which individuals and colonies of many
species potentially compete. However, substratum is not as simple a
resource as one might expect. Even solid rock surfaces are usually dissected
by small to large crevices, have different orientations with respect to each
other and to water flow, and are often made of varied rock types. Soft
substrata can be extremely diverse, consisting of both organic and inorganic
materials differing in grain sizes and in chemical composition. These
materials are further modified through the activities of the residents
themselves, adding organic material, aggregating particles and sometimes
changing the nature of the substratum so much that other species can be
totally excluded. On hard substrata, biogenic effects can also be extremely
important. These include the provision of secondary substratum through
growth processes and calcium carbonate deposition, and the addition of
physical relief that affects water flow and thus the transport of resources and
propagules within the community.
The effects of habitat complexity in benthic systems are many. Here,
habitat complexity is defined as consisting of two components, (1) habitat
heterogeneity, or patchiness of habitat types across a landscape, and (2)
habitat structure, the physical or architectural component of complexity.
Thus, a complex landscape may consist of several habitat types, each with its
own degree of physical structure. One example would be a rocky intertidal
community arrayed across a variety of habitats differing in wave exposure,
desiccation stress, substratum angle and other physical parameters, but
which also contains several degrees of physical (architectural) complexity
within each habitat or microhabitat type. The middle intertidal zone, for
example, might consist of a sloping rock bench or it could be an array of
horizontal surfaces broken by numerous gullies or pools. Physically, the

S. S. Bell et al. (eds.), Habitat Structure

© Chapman and Hall 1991
212 Habitat structure and community dynamics
latter habitat is more complex than the former. Finally, the presence of
mussel beds or algal turfs provides another type of physical complexity or
'habitat structure' for organisms living on their surfaces or within the
interstices between mussels or algae.
A given landscape can be thought of as existing within several gradients
that affect habitat conditions such as depth, wave action, irradiance,
desiccation potential, and many others (Figure 11.1). Some of these factors
covary and others are reasonably independent. Thus, even if our conceptual
benthic environment were a large sheet of unbroken rock (think of a cement
jetty), there would still be at least two major gradients, distance from open
water (thus wave energy), and vertical position or depth. The latter gradient
has many covarying variables including irradiance, temperature stress,
immersion time, and susceptibility to marine and/or to terrestrial predators.
This unbroken rock surface already has several distinct habitat types created
solely by position along either gradient. If we now consider a series of such
rock surfaces (or jetties) in an estuarine region, then we have a third major
gradient to consider, that of salinity. As the scale increases, latitudinal and
other major geographic considerations add one or two more gradients to this
array of habitats and microhabitats. Three or more gradients, with several
covarying parameters, certainly provide enough heterogeneity for many
species to coexist by 'niche specialization' alone (Horn and MacArthur,
1972; DeAngelis and Waterhouse, 1987). On top of this level of hetero-
geneity, can be added the patchiness created by temporal variation and by
disturbance events within anyone habitat type. This results in an array of
patch types differing in 'successional' state and therefore in the types of
organisms occupying them. The existence of a given disturbance regime
alone provides for the coexistence or many more species via the 'fugitive-
dominant' model (Armstrong, 1976) extended to many species differing in
recruitment and competitive abilities (see models in: Yodzis, 1978; Hast-
ings, 1980; Paine and Levin, 1981; DeAngelis and Waterhouse, 1987;
Sebens, 1988). All the processes above contribute to 'habitat heterogeneity'
even before considering the effects of physical or architectural diversity
among habitats.
In addition to the effects of gradients and of disturbance on this model
benthic system, there are at least two 'architectural' phenomena of major
importance to the diversity of species within a community and to the
coexistence of particular species. The first of these is the physical complexity
of the substratum itself. In our hard substratum model, consider the effect of
different rugosities (Harlin and Lindbergh, 1977; Norton and Fetter, 1981;
Norton, 1983) size and extent of crevices and larger channels, and even
mobilities of substratum (e.g. small to large boulders). Clearly the way in
which the substratum itself is shaped affects which species can persist and
how many species can coexist. Large immobile surfaces develop very
different communities than do small rocks that are frequently overturned
during storms (Osman, 1977; Sousa, 1979). Similarly, crevices provide
 Habitat structure and community dynamics 213
Landscape of habitat types

Gradient 1

Gradient 4

Within -habitat structure

Figure n.1 Hypothetical landscape showing at least two gradients, (1) horizontal
(e.g., wave action) and (2) vertical (e.g., depth, light, temperature) overlying a
patchy substratum of vertical (V), sloping and/or horizontal (H) surfaces inter-
spersed with boulders (B), loose gravel and sand (S). Note that sand may act as an
absolute barrier for certain benthic predators, and that vertical walls may present a
relative barrier for particular mobile species. Within anyone habitat patch, there is a
finer scale of patchiness resulting from disturbance, recolonization, and priority or
density-dependent effects of resident species (patterns in Figure close up). There are
also microhabitat gradients caused by substratum orientation which mayor may not
be the same as the overall environmental gradients (gradients 3, 4).

desiccation protection for certain species which can then outcompete other
species that normally live on exposed surfaces at that intertidal level.
Desiccation-resistant species devote much of their energy, and modify their
growth form, to be able to survive desiccation and other physical stresses on
exposed surfaces.
The second 'architectural' phenomenon is provided by organisms that
come to inhabit each type of surface. Large creatures with hard shells
(mussels, oysters, barnacles) provide secondary substrata on which other
species settle and grow, and provide sheltered microhabitats in which
certain species find refuge. One might expect the more complex, large and
stable the biogenic structure, the greater its likely effect on species diversity
and on the types of species that can exist in that habitat (e.g. Suchanek,
1979). On coral reefs, however, plating corals provide broad undersurfaces
that become colonized by many sponge, bryozoan, and ascidian species
(Buss and Jackson, 1979). Branching corals usually have polyps on all sides
of each branch, and thus provide few areas for such colonists, and mounding
214 Habitat structure and community dynamics
corals often have few accessible undersurfaces relative to plating corals
which generate large flat undersurfaces without living tissue covering them.
Thus, whereas branching corals could be considered the most complex
architecturally, and mounding corals the least, it is the plating corals of
'intermediate' architectural complexity that happen to provide the best
habitat for this cryptic community of reef organisms. The form and biological
characteristics (e.g., where the living tissue stops) of biogenically important
species may be more important than any measurable index of physical
complexity (e.g., surface to volume ratio).
The following discussion outlines several examples, from marine benthic
systems, in which the physical structure of habitat plays a major role in
determining the characteristic assemblage of species that can coexist. It has
been amply demonstrated, in virtually all ecosystem types, that the number
of habitat types, microhabitats, or potential 'niches' affects species'
coexistence and diversity (Huston, 1979). Clearly, the more different types
of habitat that exist, the higher diversity can be. Similarly, it is now
abundantly clear that the disturbance regime (i.e., space clearing provision
of primary substratum) is a prime factor determining both species diversity
and particular examples of species exclusion or coexistence (Dayton, 1971,
1984; Levin and Paine, 1974; Connell, 1978; Connell and Sousa, 1983;
Sousa, 1979, 1984; Menge and Lubchenco, 1981; Paine and Levin, 1981).
Therefore, although these effects will be discussed where needed, I will
concentrate on the effects of physical (architectural) complexity which will
be referred to as 'habitat structure'. Is there a special phenomenon or
process here, or does added habitat structure simply mean more micro-
habitats to be occupied? Three examples will be discussed, (1) substratum
physical complexity in rocky subtidal benthic communities, (2) mussel beds
in intertidal and subtidal benthic habitats, and (3) biogenic structures in soft
sediment communities. Although there are very many additional examples
of habitat structure and its effect on benthic communities, these examples
serve to illustrate the important processes and have received a good deal of
recent attention by community ecologists working in benthic marine
systems.

11.1 EXAMPLES FROM MARINE BENTHIC SYSTEMS

11.1.1 Substratum complexity in the rocky subtidal zone

Rocky coasts offer a wide range of habitat types in both the intertidal and
subtidal zones. One of the most noticeable 'architectural' features of such
coasts is the degree to which the rock itself is broken up into benches, plates,
crevices, gullies, boulder zones, pinnacles, and so on. This can be seen very
easily in the rocky intertidal zone at low tide, and the effect of the different
substratum types and orientations has long been recognized (Lewis, 1964).
Many of the differences in species assemblages on different substratum
 Examples from marine benthic systems 215
types and inclinations in the intertidal zone result from the different degrees
of protection offered against stresses such as desiccation and wave-induced
dislodgment (Dayton, 1971; Denny et al., 1985). In the shallow subtidal
zone (1-20 m depth), the same variety of substratum architecture (Figure
11.2) occurs and, predictably, the assemblages of species occupying the
different surfaces are characteristic (Connell and Keough, 1985). Large flat
areas of bedrock typically support algal assemblages including mixed turfs
and kelp beds. Where urchins or other grazers are abundant, such habitats
are characterized by crustose coralline algae and a few other grazer-resistant
species, including large subtidal mussels (Witman, 1985) and erect sponges
(Sebens, 1985). Such areas are frequently colonized by ephemeral fleshy
algae and by various sessile invertebrates, but these are consistently
removed by omnivorous grazers such as sea urchins or, in the case of certain
invertebrates, by bulldozing activities of herbivores (Dayton, 1971).

(a)

(b)

Figure 11.2 Habitat structure provided by the physical architecture of the

substratum in the rocky subtidal zone (e.g., rock walls, channels, boulders). Note
that here and in the examples in Figures 11.3 and 11.4, additional physical 'structure'
(a to c) provides diverse microhabitats as well as protection from certain physical
stresses and, potentially, refuges from particular predators.
216 Habitat structure and community dynamics
Where grazers are unable to prevent the establishment of foliose
canopy algae, such algae frequently overgrow sessile invertebrates and
crustose algae (Dayton, 1975; Lubchenco, 1978; Witman, 1985). The
development of an algal canopy can reduce water flow to the substratum,
and thus may also limit food availability to sessile invertebrates below the
canopy. Also, the 'whiplash' effect of algal fronds in wave-generated surge
prevents the establishment, or hinders the growth, of other algal species and
of many sessile invertebrates (Dayton, 1975). Where a kelp bed develops, or
where stands of other large algae form, a multilayered architecture
develops. This affects the type of epibionts that can exist on the canopy
algae, and the type and number of mobile invertebrates using the algal
fronds as microhabitats (Dahl, 1973; Dean and Connell, 1987). The fronds
provide a food resource for herbivores, such as small gastropods and
amphipods, and produce an understorey environment conducive to the
growth of low-light adapted algal species (obligate or facultative under-
storey, Dayton, 1975) and which might not otherwise be able to inhabit that
surface.
Even a crustose coralline algal assemblage provides some 'architecture'
that can be important. For example, the thick and lumpy coralline
Lithothamnion glaciale, in the Gulf of Maine, provides substratum for
boring polychaetes whereas the thinner and smoother Phymatolithon
rugulosum does not; the latter species is predictably overgrown by
Lithothamnion. Urchin grazing breaks off edge pieces from the former
species, and usually not from the latter, although both species are grazed to a
similar extent (Sebens, 1986). The long-term coexistence of the two algae
species, which often make up over 80% of the substratum cover, thus
depends on the biogenic architecture provided by one of the two competing
species, which provides a microhabitat for one or more polychaete
species.
Vertical or undercut surfaces in the shallow subtidal zone support an
entirely different assemblage, usually dominated by encrusting sessile
invertebrates and by a few low-light adapted algae (Sebens, 1985, 1988;
Vance, 1979). However, where sea urchins are very abundant, even vertical
surfaces can be dominated by crustose coralline algae (Sebens, 1986).
Vertical surfaces offer several conditions conducive to the development of
an invertebrate community: low light levels, reduced sediment loading,
water flow enhancement or disruption, and reduced access for certain
predators, such as urchins. Vertical surfaces, besides providing the correct
environment for sessile invertebrate growth, are also fragmented in space,
thus acting as habitat 'islands' , analogous to mountain tops or ponds. In the
Gulf of Maine subtidal zone, vertical rock wall communities are dominated
by species whose larvae have very short-range dispersal (Sebens, 1983, 1985;
Keough, 1983), of the order of centimetres to tens or hundreds of metres.
Very few species have the long-distance planktotrophic larvae characteristic
of intertidal invertebrate species (Dayton, 1971; Paine and Levin, 1981;
 Examples from marine benthic systems 217
Wethey, 1984; Gaines and Roughgarden, 1987). Therefore, vertical rock
walls separated by sand channels or by expanses of horizontal or sloping
rock are somewhat independent in their population and community
dynamics, and many of the species are likely to have recruitment rates that
depend on the density of adults on the same or adjacent rock walls.
Therefore, in addition to the physical structure provided by irregular rock
structures, and the heterogeneity provided by physical gradients and
disturbance, there is a degree of isolation and separation conferred by
physical structure as well. In such cases, physical or architectural complexity
provides the opportunity for chance events to become extremely important
on isolated pieces of substratum. Jackson (1977) noted that competitive
interactions are likely to be very different on small separated substrata than
on large contiguous pieces because of the randomness associated with
recruitment and later encounter probabilities (see also Keough, 1983;
Connell and Keough, 1985). Sutherland (1974, 1981), for example, gives
numerous examples of unpredictable and entirely different assemblages
developing on small isolated settling panels at the same depth.
In rocky subtidal communities along the coast of northern Massachusetts,
vertical rock walls are isolated enough that the population dynamics of
species and the types of assemblages that develop on walls only metres apart
can be quite distinct (Sebens, 1985, 1986). A series of events in 1983-1985
illustrates the semi-independent nature of these rock wall assemblages.
Three primary study sites were established in 1978 for monthly photographic
monitoring: Halfway Rock, Shag Rocks, and Inner East Point. Population
densities of mobile species and percentage cover and counts of sessile
species were calculated from photographic quadrats over this period. In
1983 a nudibranch, Tritonia plebeia, appeared and its populations increased
over 1984 until it was the most abundant nudibranch at these sites, with over
50 individuals per square metre on some rock walls (Allmon and Sebens,
1988). This specialist predator on octocorals is a northern European species
that had never been recorded in the northwest Atlantic region previously.
The time course of its population increase differed at sites just tens to
hundreds of metres apart such that, at two of the sites, they lagged about a
year behind the initial site where the species was observed. The third site,
Halfway Rock, never supported a population of this species, although this
site experienced another strong perturbation; a unique high temperature
event in 1984 that did not occur at the other two sites. Finally, all sites except
Inner East Point had dense urchin populations throughout the study period
but experienced a dramatic increase, approximately doubling adult densi-
ties, in 1984-1985. This resulted in frequent urchin excursions onto vertical
walls with subsequent removal of most invertebrates by this omnivorous
predator.
The results of these three perturbations or 'natural experiments' were as
follows (Table 11.1). At the site without urchins but which did have
numerous Tritonia, soft corals were reduced to less than a quarter of their
218 Habitat structure and community dynamics
Table 11.1 Results of three infrequent perturbations on isolated subtidal rock wall
assemblages (Allmon and Sebens, 1988; Sebens MS).

Site assemblage TR1* UR TOC Changes

1978""'--+ 1988
Inner East Pt. Invert. 0 0 0 Minor
Inner East Pt. Alcyonium Y 0 0 Alcy. (-) Recovery
Inner East Pt. Alcyonium IT Y 0 0 Alcy. (-) Recovery
Shags Outer Alcyonium Y Y 0 Alcy. (-) Met. (+)
Shags Outer Metridium Y Y 0 Minor
Shags Inner Invert. Y Y 0 Invert. (-) Crusts (+)
Shags Inner Coralline 0 Y 0 Minor
Half.Rk Outer Invert. 0 Y Y Invert. (-) Crusts (+)
Half.Rk Inner Coralline 0 Y Y Minor
Half.Rk Inner Invert. 0 Y Y Invert. (-) Crusts (+)

TRI* , arrival of an exotic nudibranch species (Tritonia plebeia) , a specialist predator

on the soft corals, Alcyonium siderium. UR, doubling of adult urchin population and
subsequent increase in urchin foraging on vertical surfaces. rc, single fall period of
abnormally high temperatures at Halfway Rock study site, which did not occur at other
sites studied. Y, event occurred, 0, event did not occur, (-), large decrease, (+), large
increase in percentage cover. Crusts, coralline algae and fleshy red crustose algae.
Recovery, partial return to original condition since 1985.

initial density in two years, after which the nudibranch disappeared and the
populations of soft corals began to recover. Other invertebrates were not
affected and remained at high percentage cover. At the Shag Rocks site,
with increases in both Tritonia and urchin density, the soft coral disappeared
together with most of the encrusting invertebrate community; this condition
has persisted to date (1988) even though Tritonia has disappeared since
1986. Finally, the Halfway Rock site, with elevated early autumn tempera-
tures in 1984, an urchin increase, but without Tritonia showed reduction of
the Alcyonium and Metridium populations initially, before the urchin
increase, followed by removal of most of the invertebrates from the walls.
Because the urchin populations have remained high for the past several
years, the walls at the Shag Rocks and at Halfway Rock have remained
dominated by crustose algae from 1985 to 1988. At both Shag Rocks and
Halfway Rock, walls which were originally dominated by crustose coralline
algae remained that way throughout this period. These are walls with easy
urchin access and frequent grazing, preventing an invertebrate community
from developing even when urchins were at the lower densities. The
urchin/coralline assemblage is thus extremely stable and showed little
change even with a doubling of the urchin population.
This series of events illustrates several important effects of habitat
structure. First, added physical complexity provides refuges and barriers.
 Examples from marine benthic systems 219
Rock walls and undercut surfaces are difficult for certain predators,
primarily urchins, to forage on in heavy wave surge and thus a refuge exists
(Laur et al., 1986). Second, added physical complexity provides numerous
new microhabitats. Our observations indicate that species are zoned on indi-
vidual walls with the largest suspension feeders, the anemone Metridium
senile, at the top of the walls and other species below. Passive suspension
feeders become uncommon at the bottom of walls, which are areas of
reduced water movement. Third, added physical complexity, which includes
separated units of substratum, increases the stochastic element of com-
munity structure. The predator Tritonia colonized one rock wall area in
1983, two nearby areas a year later, and Halfway Rock, about 7 km distant,
was never colonized although sites even farther away were. Also, the
complexities of coastal water flow are such that one site can experience
maximum temperatures several degrees higher than a site just a few
kilometres away in one year but not in others (Allmon and Sebens, 1988).
Random factors can affect recruitment, persistence, mortality, and other
demographic parameters of species on habitat 'islands'. The more the region
is fragmented, with barriers between units, the more important such factors
become and the more heterogeneous the resulting assemblages will be when
many such units are combined.

11.1.2 Mussel beds in intertidal and subtidal habitats

Mussels are one of the most ubiquitous organisms providing biogenic
structure in rocky intertidal and subtidal communities. Their relatively large
size, hard surfaces, and long-term persistence make them the 'trees' of the
mid-intertidal zone where they are usually able to outcompete macro algae
(Paine and Levin, 1981). In subtidal habitats where intense urchin grazing,
or lack of light, prevents macro algal growth, mussels form large long-lived
aggregations that provide secondary substrata for plant and animal epi-
bionts (Figure 11.3). Their interstices provide a refuge for mobile species
that would normally suffer intense predation (Witman, 1985). In the
mid-intertidal zone, mussel beds provide secondary substrata and a refuge
for a diverse assemblage of infauna which could not survive the intense
physical stresses without such protection (Suchanek, 1979). Although
mussels usually dominate this zone, in a very few localities, other groups
such as large ascidians take their place and provide comparable physical
structure (Suchanek and Paine, 1983).
The physical complexity of a mussel bed results from the number,
maximum sizes, and size distribution of mussels. These factors are, in turn, a
result of intertidal level and the history of a particular 'patch' of the
mid-intertidal zone. After a disturbance which removes a piece of mussel
bed, the newly cleared substratum is recolonized, first by non-mussel sessile
biota and then by small mussels forming a monolayer, some of which later
grow to full adult size. A mature high intertidal bed remains a monolayer of
220 Habitat structure and community dynamics
(a)

(b)

(e)
-«.
~ -
\
. ,"
. • • • •

.
N.
.,
,...:. it.~
~
1 \

Figure 11.3 Habitat structure provided by biogenic structures, the shells of

bed-forming mussels. Note that the surfaces of the shells provide secondary substrata
for attachment, and the interstices offer refugia and collect sediment, providing a
variety of microhabitats. Larger mussels and thicker beds (a to c) increase structure.

small mussels whereas a mature low-intertidal bed has a wide range of

mussel sizes and several layers of mussels attached to each other, with young
individuals mixed in (Suchanek, 1979, 1981; Paine and Levin, 1981). Such
beds also capture a sediment layer at their base, providing a type of
microhabitat, for deposit feeders, not normally found on solid rock. The
number and diversity of infaunal species thus increases dramatically with
habitat structure, providing for a much more diverse community at the
mid-intertidal level than could exist without the mussel matrix (Suchanek,
1979).
Mussel beds also occur on isolated units in some locations, or in
continuous bands in others. Because they provide a refuge for small mobile
species, they affect the community structure on adjacent surfaces as well.
Intertidal mussel beds, for example, provide refuge for limpets and other
small grazing invertebrates which forage on the mussel surfaces and on the
rock nearby (Dayton, 1973; Suchanek, 1979; Sousa, 1984). They move out
 Examples from marine benthic systems 221
of the mussel bed matrix with the rising tide and retreat as the water recedes,
thus producing a heavily grazed 'halo' of rock around individual mussel beds
and around the edges of large 'patches' within contiguous mussel beds, when
such patches are at early 'successional' stages and are dominated by
macroalgae.
Subtidal mussel beds also offer protection to grazers, particularly juvenile
sea urchins (Witman, 1985). These small urchins, and the large adults
foraging outside and on top of the mussel beds, keep the mussel surfaces
cleared of macroalgae and other fouling organisms. Witman (1987) demons-
trated experimentally that mussel beds with few urchins develop a kelp
cover and that storms dislodge mussels with attached kelp more readily than
mussels free of kelp. Although both mussels and urchins can exist without
each other, this interaction can be considered a 'facultative mutualism' in
which mussels offer a refuge from predation and urchins enhance the
survival probability of the mussels thus enhancing substratum physical
complexity. If the area were to become a kelp bed, another assemblage of
sessile epibionts might prosper, as well as fish and certain mobile inverte-
brates that utilize the kelp blades and holdfasts as refugia (reviewed by
Dayton, 1985; Ebeling and Laur, 1988; Duggins, 1988). Suchanek (1979)
suggested a similar facultative mutualism for intertidal mussels and molluscan
grazers.
Mussel beds thus affect community structure in a number of ways. First,
they increase physical structure and provide numerous microhabitat types
which can support species adapted to those conditions. Second, they provide
refugia from predation and from physical stress, favouring mobile species
which can find prey within or adjacent to the beds. Third, because mussel
beds are subject to exfoliation during storms, their particular biology and
mode of attachment provides for a successional cycle in the mid-intertidal
zone whereby patches are cleared, occupied by algae, barnacles and other
groups for several years, then gradually develop into full mussel beds again.
The fact that mussel beds also occur as isolated units probably means that
'island effects' are important here as well, although this aspect of mussel bed
dynamics remains to be explored.

11.1.3 Biogenic structures in soft sediments

Soft sediments are characterized by relatively low structural complexity,
except at the level of grain size, type and aggregation (Figure 11.4). Much of
the structure that can be identified in these habitats results from the activities
of burrowing organisms and their predators. A quick examination of a tidal
sand or mud flat reveals numerous holes, some surrounded by mounds of
excavated sand and castings defaecated by burrowing polychaetes. Often,
there will also be small to large pits or depressions caused by predators
digging into the substratum to remove their more sessile prey (Virnstein,
1977; Reise, 1985). Such small-scale disturbances of the sediment surface
222 Habitat structure and community dynamics
(a)

(b)

(e)

I ... ~I'._...._,.......
......
... '~

Figure 11.4 Habitat structure provided by biogenic structures, the burrows,

castings, and tubes of soft·substratum macrofauna (polychaetes, bivalves). As in
previous examples, additional structure (a to c) provides greater microhabitat
diversity, refugia, and attachment sites. (c) illustrates protruding tube ends of the
polychaete Diopatra with attached epifauna.

partially determine the distribution of certain infauna, including ampeliscid

amphipods, hemichordates, and harpacticoid copepods (Thistle, 1980).
The small-scale patchiness of substratum characteristics is a result
of all such processes, and of the action of bacteria on the faecal
material of deposit feeders once it has been deposited on the sediment
surface. In certain fine sediment habitats, much of the sediment is in the
form of faecal pellets (pelletized) resulting from the activities of a few
dominant polychaete species. Ecological studies have shown that the
activities of deposit-feeding species affect the sediment so as to make it less
suitable for most suspension feeders (Rhoads and Young, 1970; Wilson,
1979, 1984; Peterson, 1979; reviewed in Reise, 1985). On the other hand,
once the substratum has been bound up into more-or-Iess permanent tubes,
 Examples from marine benthic systems 223
it is less easy for the deposit feeders to continue their activities (Woodin,
1974, 1978; Gallagher et al., 1983) and if deposit feeders are active in the
sediment surface layer, larvae and juveniles of suspension feeding species
have a difficult time becoming established and maintaining their bur-rows or
tubes in the face of repeated burial and predation (Woodin, 1978, 1981).
Some polychaetes make very long-lasting and intricate tubular structures,
such as the sand-grain tubes of Pectinaria, the compacted sediment tubes of
many species, and the parchment-like tubes of Diopatra, often with attached
debris on their upper ends. Diopatra tubes act as attachment sites for
macro algae and the tube-caps provide secondary substrata for an assemb-
lage of amphipods and copepods (Bell and Coen, 1982)_ Such tubular
structures organize the soft sediment habitat in such a way that small deposit
feeding species can forage in the sediments between tubes, where they are
also relatively free from the predatory activities of pit-making predators
such as horseshoe crabs, blue crabs, and skates (Woodin, 1978, 1981; Bell
and Woodin, 1984; Brenchley, 1976, 1981). Although these structures
provide a refuge from predation for certain species, they may also prevent
the establishment of populations of certain other large deposit feeders and
other suspension feeders (Gallagher et al., 1983). In this way, tube matrices
may act similarly to plant roots in seagrass habitats (see Chapter 14). Not all
tube-building species can prevent this type of predation. For example, tube
dwelling ampeliscid amphipods are heavily preyed on by grey whales which
form large pits in subtidal soft substratum habitats (Oliver et al., 1983;
Nerini and Oliver, 1983).
The presence of large projecting ends on tubes of Diopatra also affects
water movement near the sediment surface. This has important con-
sequences for larval recruitment and sediment resuspension. Eckman et al.
(1981) have shown that flow profiles above arrays of tubes can be very
irregular and that flow can actually be higher at certain levels of the tube
matrix than in the free-stream flow above the bottom. Some tubular
structures, depending on size and density, could thus cause added sedi-
mentation and larval retention, whereas other structures might have just the
opposite effect, causing scouring of the substratum surface and making
settlement of certain larvae difficult (Eckman, 1979, 1983).
Within the sediment, gradients of water movement and oxygen con-
centration are also important. The grain size composition of sediment
affects these processes, as does the presence of tubes or roots. Often, there is
an anoxic layer a few millimetres to many centimetres below the sediment
surface, and this layer is deeper around burrows and roots where oxygen
enters the sediment through these structures. The activity of burrowing
deposit feeders affects the depth of this bioturbated layer and thus the
composition of the surface sediments (Rhoads et al., 1978). The presence of
physical structuring elements within this layer has analogous consequences
on both the particle spectrum and on layering within the sediment.
Therefore, microhabitat diversity is increased by the presence of structures
224 Habitat structure and community dynamics
both because they provide a refuge, and because they alter the physical and
chemical gradients within the sediment itself (Aller, 1982; Reise, 1987;
Meyers et al., 1987). An increase in diversity within structured sediments is
thus to be expected.
Compared to mussel beds, tubular structures and permanent burrows
within soft sediments have similar effects on the associated community.
However, for the latter structures it is primarily through their modification
and amplification of microhabitat characteristics rather than because they
act as secondary substratum, although this too occurs in certain cases (e.g.,
Diopatra, Bell and Coen, 1982). Because mussels and other hard-shelled
species provide a secondary substratum, they have an additional diversifying
effect on the associated community. Disturbance by wave action during
storms clears patches within mussel beds and, in soft sediment communities,
results in resuspension of the surface layer if that layer is not bound up into
large aggregates or protected by a dense tubular matrix. The overall effect of
such structures on disturbance to the associated community may be a
preventative one, whereby both physical disturbance and patch-forming
predatory activities are reduced when dense tubular matrices are present
(Woodin, 1976). Mussel beds, by contrast, become more susceptible to
complete removal by storms as they age because they lose their direct
attachment to the rock and become filled with empty mussel shells and
sediment (Paine and Levin, 1981). It is not clear that tubular matrices in soft
sediment suffer a similar fate, and thus their refuge function may be more
persistent.

11.2 PROCESSES CONNECTING HABITAT STRUCTURE

AND COMMUNITY DYNAMICS
The previous section examined several examples in which increased physical
complexity increases diversity or allows coexistence of particular species
such as predators and prey or competitors for space. It was noted that there
are several different processes whereby additional physical structure acts on
components of the community or local assemblage. In this section, several of
these processes will be examined in some detail as general concepts related
to the role of habitat structure.

1.2.1 Habitat or microhabitat diversity and area relationships

The most obvious effects of increased physical or architectural complexity
are increased surface area and the resulting potential for increased
microhabitat diversity. Greater surface area provides more attachment,
perch, or refuge space for more individuals of a given species and thus the
potential for coexistence of several species if such coexistence is governed by
external factors such as sources of mortality or limitations on recruitment.
Added surface area alone would not be expected to increase diversity or to
 Habitat structure and community dynamics 225
allow long-term stable coexistence if two species could not otherwise
coexist; it would simply provide more area for the competitive dominant.
Some mechanism such as the 'fugitive-dominant' model (see models in
Armstrong, 1976; DeAngelis and Waterhouse, 1987; Sebens, 1988) must act
to permit coexistence of competing species on an otherwise homogeneous
surface before added surface area can act to enhance this result. More
surface area then reduces the chance that the rarer of the two species would
be displaced on a given piece of substratum by chance events alone. In
Armstrong's (1976) model, stable coexistence between two species com-
peting for space results when disturbance rate is within a certain range, the
competitive (dominant) overgrows the inferior (fugitive) species, but the
latter species has the higher recruitment rate.
Microhabitat diversity results from the effect of added vertical relief
caused by a complex physical structure. Immediately, there is a gradient
within the structure, such as a reduction of water movement from top to
bottom and a concomitant increase in sedimentation rate near the bottom of
the structure. Certain types of physical structure enhance turbulence or
actually increase water flow rates in some part of the structure, thus
providing areas of low, intermediate, and strong turbulent flow across or
within the structure. Light levels, temperature, desiccation stress and other
important parameters are also likely to vary within such a structure. Mussel
beds or tufts of algae in the intertidal zone are good examples. Epibionts
living on upper valve surfaces of mussels experience temperatures at least as
high as those on open rock surfaces, whereas those even a few centimetres
below will experience greatly ameliorated conditions, and those near the base
of the bed may see little if any of such stressful conditions during anyone tidal
exposure. Increased microhabitat diversity with increased physical structure
is probably the most universal and important of the processes enhancing
diversity in most systems, especially those where the scale is relatively large
and where the biogenic structures or the substratum itself provide more
attachment sites with added complexity.

11.2.2 Refuges from predation and other sources of mortality

Aside from microhabitat diversification, the protection provided by com-
plex physical structures is probably the other most universal process
whereby such structures increase diversity or allow coexistence (Huffaker,
1958). This is evident in studies from many divergent ecosystems, and is
particularly important where predators are large and possibly generalized in
their diet. In such cases, physical structures whose interstices are small
relative to predator body size confer excellent refuges for one or more
species of prey (Bell and Woodin, 1984; Witman, 1985). However, there is
the potential for such structures to harbour predators as well as prey.
Suchanek (1979) noted the abundance of Nucella spp., predatory gastro-
pods, in mussel beds with large interstices and noted that they preyed
226 Habitat structure and community dynamics
heavily on small mussels within the bed. Heck and Crowder (Chapter 14)
found that seagrass communities also offer refuge to predators of inter-
mediate size whose prey are therefore more likely to suffer predation in the
seagrass bed than outside it. Scale, of both habitat and organisms, is thus very
important when considering the refuge value of particular habitat structur-
ing elements.
Refugia are also important when extreme physical conditions are a likely
source of mortality for species which can find refuge in the matrix provided
by structuring elements. The most obvious examples of this process come
from the intertidal zone where marine animals and plants must cope with
exposures of minutes to many hours between tides. Certain species, such as
barnacles and mussels, have developed elaborate morphological, physiolo-
gical and biochemical mechanisms to resist desiccation, temperature stress,
freezing, and other extremes. Such species often survive higher in the
intertidal in crevices and pools than they do on open surfaces, thus
benefiting from the added structure of the rock substratum when it is
irregular (Lewis, 1964). Even barnacle aggregations provide some protec-
tion for associated species, as evidenced by the presence of tiny snails and
arthropods nestled between them when the tide is out. Small grazing halos
can be seen around barnacle aggregations when they border patches of
microalgae, because the herbivorous snails forage only a few centimetres
from the edge of the aggregation. These are analogous to the halos around
mussel beds lower in the intertidal zone. The importance of habitat struc-
ture providing refuges from environmental stress and associated mortality
depends very strongly on the frequency and intensity of those stresses. In
fringe areas, such as the intertidal zone, this process is more likely to be
critical than in the subtidal zone where the effects of temperature, freezing,
and osmotic change are less likely to be so important. Also, because water
moves through a structure such as a subtidal mussel bed, it is likely to be at
the same temperature inside as outside because of the rapid heat conduction
of seawater, whereas intertidal mussel beds are filled with air when the tide is
out. Air conducts heat more slowly and thus large temperature gradients can
be maintained over even a few centimetres.

11.2.3 Separation, isolation and the island effect

Structures can be physically complex yet continuous in space, such that
propagules or adults can move between regions of the habitat unimpeded. A
good example of such a structure would be a contiguous mussel bed
stretching horizontally for many metres along the mid-intertidal zone, as
habitat for the small crustaceans and molluscs inhabiting the bed. On the
other hand, similar structures of equal complexity could include barriers
between subunits, or may exist as microhabitat islands in an otherwise
uninhabitable matrix (Huffaker, 1958; Jackson, 1977). Using the mussel bed
example, there are localities where intertidal mussel beds are discrete units
 Habitat structure and community dynamics 227
just a few metres across (Suchanek, 1979) and where movement of adults
would be difficult for the crustaceans, polychaetes and gastropods that live
within the beds. Those species with widely dispersed propagules would be
able to colonize such widely spaced beds, while species with brooded larvae
or juveniles would have a more difficult time doing so. This necessarily leads
to differences between individual beds and could even produce an overall
higher diversity when all such beds are combined than when a contiguous
area of mussel bed of equal size is considered. Boulder fields are another
excellent example of isolated substratum units developing very different
assemblages on their surfaces. Not only are the boulders separated in space,
but they also differ in size and thus in motility during storms. Assemblages
including long-lived species develop only on the larger boulders that are
almost never overturned, whereas smaller rocks are covered almost
exclusively by ephemeral algal species (Sousa, 1979). Intertidal pools are
another example of such habitat 'islands', where unique events and
individual pool history affect their species composition markedly (Dethier,
1984).
Subtidal vertical rock surfaces also frequently occur as disjunct units and
most of the species on shallow rock walls have some way to colonize the
same or adjacent walls rapidly, either by the production of asexual
propagules, by the release of brooded larvae or juveniles, or simply by
lateral colony growth and fragmentation (Sebens, 1985). These characteris-
tics allow certain species to monopolize most of the space on a given wall,
and probably to retard or prevent colonization by competitors, as also
occurs on settling panels, which are experimentally produced isolated units
(Sutherland, 1974, 1981). Once an encrusting species has taken over an
isolated piece of substratum, or even a large portion of one, it may be very
difficult for even superior competitors to invade and take that space away
(the priority effect). Predators may also have a difficult time locating a
patchy prey resource. Therefore, the separation of such substrata enhances
species coexistences in the larger community, and potentially increases the
overall diversity of the encrusting community over the broader area. The
stochastic nature of successful recruitment, survival, and long-term persist-
ence on small isolated pieces of substratum ensures that rock surfaces with
similar physical attributes will come to support very different assemblages at
the same time. These effects will be particularly strong in cases where
propagules have short dispersal distances and, therefore, local assemblage
dynamics are independent of a large 'pool' of potential colonists which is
essentially identical for all parts of the substratum.

11.2.4 Positive density-dependence and population dynamics

Many of the species that add to the physical structure of a habitat, and some
of those that benefit from added structure, show positive density-dependent
population dynamics. Below some minimum population density, individual
228 Habitat structure and community dynamics
fitness suffers from reproductive failure, interspecific competition, or
intense predation. Above this density, and possibly increasing with density,
individuals benefit from the presence of neighbours in a number of ways. In
barnacles, certain species attain higher fecundities when crowded (Wethey,
1984), whereas other species show the opposite trend. Those that form
dense continuous beds with attached walls benefit from certain levels of
crowding. It may be less costly to build their shell and maintain their
position, in the face of intense wave action, when sharing walls than it would
be if they were isolated. Barnacles also depend on the nearness of
neighbours to achieve internal fertilization, and thus suffer low fecundity or
are forced to self-fertilize at extremely low densities. Mussels survive better
when in beds with many neighbours than if they were to occur as isolated
individuals, a condition which is, in fact, very difficult to observe.
Contiguous beds prevent the development of strong shear forces which can
occur at edges of beds or on individual mussels. Any factors reducing the
surface area exposed to oncoming waves also reduce the probability of
dislodgement and death for intertidal invertebrates (Denny et al., 1985).
The octocoral A/cyanium siderium in the temperate rocky subtidal zone
also does better when relatively crowded, at least at juvenile stages (Sebens,
1983). Juveniles approximately 1.5-3.5 cm from adult colonies had the
highest survival rates and growth rates compared to juveniles either closer or
farther from large colonies. This is probably a result of reduced water flow,
or reduced prey abundance, near large colonies and of reduced sea urchin
grazing at the intermediate distance compared to distances greater than 3.5
cm away. Urchins tend to graze in partially cleared patches on vertical walls
and to avoid stands of large soft-bodied invertebrates such as anemones and
soft corals. Although they can consume such prey, they do so from a position
where they can maintain attachment with their tube feet even in strong wave
surge. Therefore, Alcyanium tends to occur in dense aggregations on certain
rock walls but to be absent altogether from other walls, especially where
either the anemone Metridium senile or the ascidian Aplidium canstellatum
is abundant. There is some evidence that both the latter species also have
some degree of positive density-dependence to their population dynamics,
and thus once a large part of the rock surface has become occupied by any of
these three species, they can probably prevent the others from becoming
established, or at least slow the rate at which displacement occurs (Sebens,
1985).
Positive density-dependence thus creates patches where particular species
do well because of historical events. Given a particularly good recruitment
event, or a period with low predation, a population of one species can
become dense enough to achieve added protection from predators (Sebens,
1983) or competitors (Peterson and Black, 1988). Highsmith (1982) also
found that juvenile survival was higher in beds of adults of the sand dollar
Dendraster and that larvae were induced to settle in those beds. As an
example of the effect of density on competition, Buss (1979) showed that
 Habitat structure and community dynamics 229
certain erect bryozoans were less likely to be overgrown by competitors
when they occurred in dense patches of conspecifics; this was advantageous
to individual colonies even though growth rates decreased at high density.
Wilson (1983) found a similar result for infaunal polychaetes. The result of
positive density-dependence is to allow patches of a single species, that have
developed through chance events, to persist for long periods in the face of
disturbance, predation, or competition. Again, as in the case of separated
substrata, this process can increase the number of coexisting species and
prevent competitive exclusion by a single dominant competitor, thus
increasing community-wide diversity. Isolation of substratum units makes
this process even more important, because it becomes very difficult for
competing species to invade when one colony or clone has taken over an
entire patch of available substratum (Sutherland, 1974; Jackson, 1977).
Positive density dependence is tied to increased physical structure of the
habitat in several ways. If such increased structure includes patches of
suitable substratum separated in space, then the arguments above hold true.
Even if most substratum is contiguous and there are few barriers or
separations, patches dominated by species showing positive density-de-
pendence are likely to persist in the face of advancing competitors. The
greater the physical complexity, or habitat structure, the more places there
will be where dense patches of a single species can become established.
These places need not be complete 'islands' for this phenomenon to be an
important contributor to long-term species coexistence and thus to overall
diversity.

11.2.5 Disturbance, succession, and scale effects

In this discussion of habitat structure and its effects on community dynamics,
the concept of disturbance has been invoked a number of times. However,
this has mostly been in the context of habitat structure preventing
disturbance or lessening its impact. Disturbance may, on the other hand,
have a major role in creating habitat structure over the long term. If
consideration is restricted to disturbances that clear space by causing the
mortality of sessile species holding primary substratum, then it is clear that
disturbance starts the 'successional' cycle in benthic systems (Dayton, 1971;
Sousa, 1979, 1985; Paine and Levin, 1981). Succession, in this case, refers to
a relatively predictable replacement series of organisms following space
clearing, rather than to any necessary alteration of the environment needed
for the rest of the sequence to occur (Connell and Slatyer, 1977; Chapter 8).
In North American west coast rocky intertidal habitats, disturbance in the
mid-intertidal zone clears away patches of mussel bed or clears off any other
species that colonize more rapidly than mussels. Cleared patches are
colonized first by diatoms and microalgae, then by ephemeral macro algae
and barnacles, followed by more persistent macro algae and larger barna-
cles, and finally by two species of mussels, the larger of which takes longer to
230 Habitat structure and community dynamics
establish itself. This species, Mytilus californianus, is the primary bed-
former that makes the very large and spatially intricate beds of the mid to
low intertidal zone (Dayton, 1971; Paine and Levin, 1981; Suchanek, 1981).
Habitat structure thus increases almost linearly with time since the
disturbance occurred, or with successional stage. As larger-bodied species
take over a patch, there are more places for small mobile species to take
refuge until a mussel bed finally develops which itself increases in complexity
as it matures, one day to be torn away again in a storm. Disturbance thus
regulates habitat structure in the following way; low disturbance rates allow
long-term persistence of thick mussel beds with high physical structure,
whereas high disturbance rates remove most of the larger mussel beds and
keep the community dominated by earlier successional stages with lower
vertical relief and less physical complexity. An intermediate rate of
disturbance will allow patches of mature mussel bed to coexist with patches
at all the earlier stages, thus maximizing overall community diversity by
combining increased habitat heterogeneity with complex habitat structure in
some portions of the landscape (see Chapter 12).
The preceding example introduces the concept of 'structural diversity', a
term that can be used to describe a system composed of some patches with
low structure and other patches with intermediate to complex physical
structure. A mud flat with patches of tube-forming worms interspersed with
loose sediment would be an example of a structurally diverse habitat, as
would mussel beds interspersed with patches of rock at earlier successional
stages. It might be argued, although it is by no means certain, that a
community arrayed over habitats showing a high degree of structural
diversity could support more species than a community arrayed over
habitats all of which have the same complex habitat structure. Habitat
heterogeneity and habitat structure are thus both important contributors to
community diversity and to species coexistence. Although it is not difficult
to see that either 'heterogeneity' or 'structure' alone can promote species
diversity, it is more interesting to consider the processes by which this
amplification takes place. Once these processes are examined one by one, it
becomes clear that it is the processes themselves that are important and
interesting and that the simple correlation of more species with more habitat
types, or more species with more complex architecture, is just the obvious
pattern that can lead to fruitful studies of mechanism.

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