166
Lisa A. LEVIN and Andrew J. GOODAY
and control trays in all Atlantic studies (Desbruy` eres
et al., 1980; Grassle and Morse-Porteous, 1987; Snelgrove et al., 1992, 1996). These experiments, along
with those in which direct enrichment of the bottom has
been achieved by placement of fish carcasses, wood,
or marine algae on the seabed (Grassle and MorsePorteous, 1987; Snelgrove et al., 1994) indicate the
importance of patchy organic inputs to the structure
of benthic communities. Sediments near experimentally
emplaced wood blocks yielded high densities of mussels (Idasola argentea) and wood borers (Xyloredo sp.),
while sediments containing Sargassum exhibited elevated densities of amphipods and several polychaetes
including Capitella spp. and Ophryotrocha sp. Patterns
of species richness, dominance, taxonomic composition
and abundance all seem to be affected by the supply
of organic matter to the seabed (Gooday and Turley,
1990; Rice and Lambshead, 1994; Grassle and Grassle,
1994). In recent decades it has become clear that
persistent heterogeneity of this supply results from
falling carcasses of fish and mammals, macroalgae and
phytodetritus, and from the interaction of accumulating
particles with biogenic structures such as depressions,
tests, mounds, tracks, fecal casts and traces, which
impose roughness on the seabed.
CONCLUSIONS
The Atlantic Ocean has, without question, played a
focal role in the development of deep-sea biology.
Recent discoveries and long-term, time-series investigations have yielded the unexpected, making it clear
that the Atlantic is a dynamic ocean whose inhabitants
experience environmental variation over a wide range
of spatial and temporal scales. The consequences of
this forcing, in terms of taxonomic makeup, population
dynamics and ecology, are clearly documented for
microbial and protozoan forms, but these consequences
are less clear for certain higher taxa. Within the
Atlantic, the continental margins are among the most
heterogeneous and biologically productive settings.
They continue to be most heavily studied because
of their proximity to population centers and their
commercial potential (e.g., exploitation of fisheries,
hydrocarbon resources). Despite this, knowledge of
species composition and basic functional attributes is
lacking for most places within the Atlantic, especially
south of the equator. Given that the Atlantic is
the best studied of the Oceans, this is true for
all of the deep sea. It is evident that the Western
and Eastern sides of the Atlantic have been studied
independently, with a few notable exceptions. Similarly,
the megabenthos, macrobenthos and meiobenthos, as
well as the various taxonomic groups of benthic fauna,
have been studied in isolation. Often the research
focus and methodologies have differed. As a result,
except for the issue of species diversity (see Stuart
et al., Chapter 10, this volume), there have been
only a few whole-basin syntheses for the Atlantic
Ocean (see Sibuet et al., 1989; Rowe et al., 1991).
Even patterns of species diversity have been addressed
primarily for the macrofauna. This should change with
increased potential for remote measurement in the deep
sea, acquisition of large-scale synoptic data, dramatic
technological advances in measurement capabilities in
situ, and the tremendous improvement in speed and
ease of communication between scientists in distant
places. As with diversity, ocean-scale considerations
of productivity, hydrodynamics, historical, geological,
and chemical influences, as well as integration among
biological components (micro, meio-, macro- and
megafaunal; procaryotic and eucaryotic; microbial, invertebrate and ichthyofaunal) will certainly yield better
understanding of the processes underlying biological
pattern in this remarkable ocean.
ACKNOWLEDGEMENTS
We are grateful to N.R. Merrett, A. Gebruk, A.L. Rice,
M.H. Thurston and C.M. Turley for reading various sections of the manuscript. A. Gebruk kindly
contributed information about the Russian literature.
Careful critiques of the manuscript were provided by
J. Gage and P. Tyler.
REFERENCES
Agassiz, A., 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer ‘Blake’ in the Gulf of Mexico, in the
Caribbean Sea, and along the Atlantic Coast of the United States,
from 1877 to 1880. Bull. Mus. Comp. Zool. Harv. Univ., 14:
1−314.
Aldred, R.G., Thurston, M.H., Rice, A.L. and Morley, D.R., 1976.
An acoustically monitored opening and closing epibenthic sledge.
Deep-Sea Res., 23: 167−174.
Aldred, R.G., Riemann-Z¨ urneck, K., Thiel, H. and Rice, A.L., 1979.
Ecological observations on the deep-sea anemone Actinoscyphia
aurelia. Oceanol. Acta, 2: 389−395.
Allen, J.A. and Sanders, H.L., 1996. The zoogeography, diversity and
origin of the deep-sea protobranch bivalves of the Atlantic: the
epilogue. Prog. Oceanogr., 38: 95−153.
Lisa A. LEVIN and Andrew J. GOODAY
and control trays in all Atlantic studies (Desbruy` eres
et al., 1980; Grassle and Morse-Porteous, 1987; Snelgrove et al., 1992, 1996). These experiments, along
with those in which direct enrichment of the bottom has
been achieved by placement of fish carcasses, wood,
or marine algae on the seabed (Grassle and MorsePorteous, 1987; Snelgrove et al., 1994) indicate the
importance of patchy organic inputs to the structure
of benthic communities. Sediments near experimentally
emplaced wood blocks yielded high densities of mussels (Idasola argentea) and wood borers (Xyloredo sp.),
while sediments containing Sargassum exhibited elevated densities of amphipods and several polychaetes
including Capitella spp. and Ophryotrocha sp. Patterns
of species richness, dominance, taxonomic composition
and abundance all seem to be affected by the supply
of organic matter to the seabed (Gooday and Turley,
1990; Rice and Lambshead, 1994; Grassle and Grassle,
1994). In recent decades it has become clear that
persistent heterogeneity of this supply results from
falling carcasses of fish and mammals, macroalgae and
phytodetritus, and from the interaction of accumulating
particles with biogenic structures such as depressions,
tests, mounds, tracks, fecal casts and traces, which
impose roughness on the seabed.
CONCLUSIONS
The Atlantic Ocean has, without question, played a
focal role in the development of deep-sea biology.
Recent discoveries and long-term, time-series investigations have yielded the unexpected, making it clear
that the Atlantic is a dynamic ocean whose inhabitants
experience environmental variation over a wide range
of spatial and temporal scales. The consequences of
this forcing, in terms of taxonomic makeup, population
dynamics and ecology, are clearly documented for
microbial and protozoan forms, but these consequences
are less clear for certain higher taxa. Within the
Atlantic, the continental margins are among the most
heterogeneous and biologically productive settings.
They continue to be most heavily studied because
of their proximity to population centers and their
commercial potential (e.g., exploitation of fisheries,
hydrocarbon resources). Despite this, knowledge of
species composition and basic functional attributes is
lacking for most places within the Atlantic, especially
south of the equator. Given that the Atlantic is
the best studied of the Oceans, this is true for
all of the deep sea. It is evident that the Western
and Eastern sides of the Atlantic have been studied
independently, with a few notable exceptions. Similarly,
the megabenthos, macrobenthos and meiobenthos, as
well as the various taxonomic groups of benthic fauna,
have been studied in isolation. Often the research
focus and methodologies have differed. As a result,
except for the issue of species diversity (see Stuart
et al., Chapter 10, this volume), there have been
only a few whole-basin syntheses for the Atlantic
Ocean (see Sibuet et al., 1989; Rowe et al., 1991).
Even patterns of species diversity have been addressed
primarily for the macrofauna. This should change with
increased potential for remote measurement in the deep
sea, acquisition of large-scale synoptic data, dramatic
technological advances in measurement capabilities in
situ, and the tremendous improvement in speed and
ease of communication between scientists in distant
places. As with diversity, ocean-scale considerations
of productivity, hydrodynamics, historical, geological,
and chemical influences, as well as integration among
biological components (micro, meio-, macro- and
megafaunal; procaryotic and eucaryotic; microbial, invertebrate and ichthyofaunal) will certainly yield better
understanding of the processes underlying biological
pattern in this remarkable ocean.
ACKNOWLEDGEMENTS
We are grateful to N.R. Merrett, A. Gebruk, A.L. Rice,
M.H. Thurston and C.M. Turley for reading various sections of the manuscript. A. Gebruk kindly
contributed information about the Russian literature.
Careful critiques of the manuscript were provided by
J. Gage and P. Tyler.
REFERENCES
Agassiz, A., 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer ‘Blake’ in the Gulf of Mexico, in the
Caribbean Sea, and along the Atlantic Coast of the United States,
from 1877 to 1880. Bull. Mus. Comp. Zool. Harv. Univ., 14:
1−314.
Aldred, R.G., Thurston, M.H., Rice, A.L. and Morley, D.R., 1976.
An acoustically monitored opening and closing epibenthic sledge.
Deep-Sea Res., 23: 167−174.
Aldred, R.G., Riemann-Z¨ urneck, K., Thiel, H. and Rice, A.L., 1979.
Ecological observations on the deep-sea anemone Actinoscyphia
aurelia. Oceanol. Acta, 2: 389−395.
Allen, J.A. and Sanders, H.L., 1996. The zoogeography, diversity and
origin of the deep-sea protobranch bivalves of the Atlantic: the
epilogue. Prog. Oceanogr., 38: 95−153.