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BOOK
REVIEWS
1974. The structure
of marine
Harvard University
Press, Camhlassachusetts.
x + 128 p. $7.95.
STEELE, J. H.
ecosystems.
bridge,
Pretiaceous foes, still drifting
by in numbers
unabated,
Were baffled
now by tactics which their
dining plans frustrated.
1Valter Garstang
The monograph provides an apt description
of
the utility of systems analysis in combining theory,
empiricism, and experiment for a study of the tactics of marine ecosystems.
Despite the evident
delay in printing, the material represents the next
exciting step beyond previous distillations
of marine ecological theory (Riley 1963; Margalef 1968)
both as a summary of the general art of marine
modeling up to the IBP/PhI
working conference
on this subject (held 4-8 October 1971 in Rome
at FAO) and as insight into the more recent research of one of the pioneers of mathematical
transfusion
in marine ecology.
This volume may
prove to be as influential
among marine ecologists
as MacArthur
and Wilson’s
( 1967 ) theoretical
treatise \vas among terrestrial
ecologists.
After disposing of ecological
shibboleths
such
as diversity and sensitivity analysis, Steele proceeds
to an interesting
discussion of the problems, assumptions, and rationale in construction
of simulation models.
The text is somewhat
terse in
places, and the reader might then benefit from
either a longer exposition or a thorough kno\vledge of the cited literature.
Despite his argulllents
for energy flow, the author explores the seasonal
dynamics
of nutrients,
phytoplankton,
and zooplankton within the North Sea (the benthos and
fish are uncoupled in this treatment)
in terms of
nitrogen and carbon because of data constraints.
Choice of these units rather than gram calories
may be more appropriate
in any case for consideration of biological amplification
of pesticides, of
alteration
of protein content, and of implications
A significant
and important efof eutrophication.
fort is expended throughout
the monograph
in
examining the structure and consequences of alternate formulations
of the model.
An appropriate
subtitle for the book might ha\.-e
been “An essay on the existence of feeding thresholds,” for, despite my own predilections
to adopt
the same artifice in t\\To-dimensional models (Walsh
in press), there is much ambiguity
in the experimental literature
on their actual presence or absense (Parsons et al. 1967; Frost 197.2; Corner et
al. 1972). As Steele suggests (p. 94) “the omission of plankton
patchiness
severely limits the
and
validity
of comparisons
with observations,”
LIMNOLOGY
AND OCEANOGRAPHY
his endorsement of thresholds may reflect a necessary parameterization
to restore spatial heterogeneit>, in his averaged, one-dimensional
model of the
Korth Sea. If one could come to grips with patchiness in a three-dimensional
model, the question
of feeding thresholds might then become a question of prey availability.
Steele’s interesting comparison of control mechanisms within terrestrial
and marine plankton systems may further reflect a basic property of patchiness as a result of the size of organisms among
marine trophic levels. Sheldon et al. (1973, p. 719)
point out that “there is no common pelagic equivalent of a small goat browsing on a large tree”
and “that at each link in the food chain large particles take up smaller ones.” Each lower trophic
level of smaller organisms may represent a patchy
resource to its predator
in the open sea. Thus
Steele’s speculation
( p. 107) that “there is no
suggestion that the energy flow at the plant-herbivore levels was significantly
changed by the alteration in the fish population,”
with rejection (also
p, 107 ) of the alternative
terrestrial
hypothesis
(Hairston
et al. 1960) “that predator control is
essential for lower trophic levels,” may reflect human ability to solve their prey patchiness problems
\\-ith acoustic gear and purse seines, while, irrespective of the removal of predation pressure, zooplankton have not yet evolved such techniques or
tactics.
In contrast, if one considers a marine system
\vith large plants of fixed location such as macrophytes, the prey patchiness problem may not be
as important
as in pelagic systems. Steele considered a macrophyte
system to be a terrestrial
analog, and recent work in St. ?Llargaret’s Bay,
Canada, suggests that the control mechanisms of
this system may, indeed, be of the terrestrial type.
5lann (1973) advanced the hypothesis that carnil-ore control may be important, with human predation on lobsters leading to overgrazed kelp beds
by sea urchins (the lobster’s prey), similar to terrestrial overgrazing by deer on the Kaibab plateau
after removal of their predators.
I noted only a few typographical
errors, the
price of the book is reasonable, and the contents
should be of interest to all environmental
scientists.
John J. Walsh
Department
of Oceanograph)
University
of \Vashington
Seattle
98195
References
CORSER, E. D. S., R. N. HEAD, AND C. C. KILVINGTON.
I.972.
On the nutrition
and me-
1015
NOVEMBER 1974, V. 19 (6)
1016
BOOK REVIEWS
tabolism of zooplankton.
8. The grazing of
Biddulphia
cells by Calanus helgolandicus.
J.
Mar. Biol. Assoc. U.K. 52: 847-861.
FROST, B. W. 1972. Effects of size and concentration of food particles on the feeding behavior of the marine planktonic
copepod Calanus
pacificus.
Limnol. Oceanogr. 17 : 805-815.
HAIRSTON, N. G., F. E. S~LIITH, AND L. B. SLOBODKIN. 1960. Community
structure, population control and competition.
Am. Nat. 94:
421-425.
MACARTHUR, R. H., AND E. 0. WILSON.
1967.
The theory of island biogeography.
Princeton Univ.
MANN, K. H. 1973. Seaweeds: their productivity and strategy for growth.
Science 182 :
975-981.
MARGALEF, R. 1968. Perspectives
in ecological
theory.
Univ. Chicago.
PARSONS, T. R., R. J. LEBRASSEUR, AND J. D.
FULTON.
1967.
Some observations
on the
dependence
of zooplankton
grazing on the
cell size and concentration
of phytoplankton
blooms.
J. 0 ceanogr. Sot. Jap. 23: 10-17.
RILEY, G. A. 1963. Theory of food-chain
relations in the ocean, p. 438-463.
In hl. N. Hill
[ed.], The sea, v. 2. Interscience.
SHELDON, R. \li., W. H. SUTCLIFFE, JR., AND A.
PR~KASH. 1973. The production
of particles
in the surface waters of the ocean with particular reference to the Sargasso Sea. Limnol. Oceanogr. 18: 719-733.
WALSH, J. J.
In press.
A spatial simulation
model
of the Peru uplvelling
ecosystem.
Deep-Sea Res.
DEGENS, E. T., AND D. A. Ross
[EDs.].
1974.
The
Black
Sea-Geology,
chemistry,
and
American Association of Petroleum
biology.
Geologists,
$33.00.
LAKING, P. N.
ology,
Woods
Woods
$10.00.
Tulsa,
1974.
chemistry,
Oklahoma.
ix +
633
p.
The
Black
Sea:
Its gebiology.
A bibliography.
Hole
Oceanographic
Institution,
Hole, Massachusetts.
xiv + 368 p.
The Black Sea has long fascinated earth scientists because of its anoxic bottom waters and low
salinities.
For English-speaking
scientists literature
on the Black Sea has previously
been difficult
to
obtain, much of it in Russian aside from a few
review papers such as Casper’s paper in the Treatise on marine ecology and paleoecology
(Casper
1957 ) .
The 45 well organized papers in this sizable
volume remedy that problem.
The papers, completed in late 1971, include results from several
expeditions to the Black Sea: RV Pillsbury of the
University
of Miami
( 1965 ), NOAA’s Oceanographer (1967), and primarily
the NSF-ONR
sponsored 7-week cruise of the RV Atlantis II of r$‘oods
Hole Oceanographic
Institution
( 1969).
Papers
and Turkish
scientists
from Russian, Bulgarian,
summarize results from the long-active
Black Sea
studies in those countries to provide access to a
literature
unfamiliar
to most American scientists.
Papers from other IVestern
European
scientists
complete the wide coverage. The companion bibliography,
covering the Black Sea literature
from
1890 to 1973, will be useful for future studies of
the area.
The Degens-Ross book has a distinctly
geological flavor.
Geological structure and history, sediments, and geochemistry
are the subject of 27 of
the 45 papers. And the nine papers on biology
emphasize marine organisms whose skeletal remains are likely to be found in sediments-molluscs, diatoms, coccoliths.
Despite the geological
focus of the volume, the geology suffers from the
2-year delay betxveen completion
of the papers
For instance, recent advances in
and publication.
the rapidly
developing
plate tectonic theory are
not fully reflected in many of the geological papers, giving them a curiously dated tone. hluch
of the geological attention is directed to\vard the
history of the past 25,000 years, with much less
attention given to the formation of the depression
that contains the Black, Caspian, and hlediterranean Seas.
Physical oceanography
received little notice.
I
think that a summary paper reviewing
the circulation of the Black Sea, regional hydrology,
and
climatology
would have been a useful addition to
the volume. Some of the needed data are included
in the various papers but are too scattered to be
very useful. In the same vein, a sur~unary article
on primary productivity
in the Black Sea could
have provided the needed framelvork
for the several papers on specialized
aspects of Black Sea
biology.
Perhaps my prejudices
are showing,
but I
thought that the papers on geochemistry
were excellent.
Because of the particular
significance
of
the Black Sea, organic geochemistry
received special attention.
And results will interest anyone
concerned
with hydrocarbon
formation
and alteration in natural \yaters and sediment deposits.
The papers on trace elements and sulfides in sediments add to our understanding
of chemical processes in lo\\‘-oxygen marine environments.
The book is nicely done. It has an attractive
layout; the figures are easily legible; and the text
is free of obvious typographic
and grammatical
errors. For that the editors are to be commended.
The abstracts for each paper and a summary chapter by K. 0. Emergy and J. hl. Hunt are quite
useful.
The computer-generated
index in the Degens-