The Structure of Marine Communities over Time: Long-
and Short- Term Changes and hydrography.
The Structure
14.1 Long- and Short-Term Changes
Conditions and organisms change on marine
environments at a wide of scales, from minutes to
thousands of years (Haury et al., 1978). Even great
depths near the bottom where temperature and the
chemical milieu are rather constant, there are pulses of
materials raining down from seasonal cycles of
production near the surface (cf. Section 10.1121).
Presumably this is why seasonal breeding takes place
even in benthic organisms in the otherwise very
constant deep sea (Grassle and Sanders, 1973; Rokop,
1974, 1977), The _magnitude of production at the
surface also varies from year to year, so that bottom
fauna are exposed to variation over years as well as
months.
The relative time scale of changes needs to be
expressed in relation to the life span of organisms.
Organisms generally respond to environmental changes
that are shorter than one generation time by using
behavioral (hiding, inactivity) or physiological
mechanisms (dormancy, resting stages, tolerance). One
of the most well-documented responses to short* term
changes is the daily vertical migration of organisms
through water or sediment columns. Although only one
of many short-term phenomena vertical migration is so
prominent that it is given separate treatment below.
Longer-term changes in conditions, in time scale of
months, years, and longer are also pervasive and
prompt significant changes in communities of
organisms. Seasonal cycles are one such kind of change,
and are dis-cussed below. Other less obviously periodic
changes such as variations in wind direction and
strength, surface temperature, and salinity (Fig. 14-1)
are also common. Other phenomena that change
structure of marine communities on different time
scales are discussed in Naylor and Hartnoll (1979). It is
difficult to identify concretely how such changes affect
marine communities, but there are pronounced
changes in the abundances of many species that are
related to large-scale, long-term variations in climate
The waters around the British Isles are among the best
and longest studied of any marine environment. During
the 1920s the cold-water, open-ocean community in
the English Channel (dominated by the chaetognath
Sagitta elegans and herring) was replaced by a warmer-
water assemblage (dominated by S. setosa and another
clupeid fish, the pilchard). Accumulations of nutrients in
surface waters during the winter were lower, spring
blooms of phytoplankton were accordingly reduced,
and zooplankton and demersal fish were reduced in
numbers. About 1965 the trends in the Western English
Channel reversed and by 1975 there were virtually no
pilchard left, and zooplankton and larvae of demersal
fish increased to the levels of the 1920s, In 1979 S.
setosa was scarce or absent and S. elegans was the
dominant chaetognath. The transition is not complete;
there are occasional short-term reversals to warmer-
water faunas. The herring have not returned in large
numbers; rather the mackerel has become the
dominant pelagic fish, The specific mechanisms behind
these major changes in the pelagic community in the
English Channel are not well understood (Southward,
1980), but there are three major points emphasize: (1)
the ocean is a continuously changing environment,
where changes can be fast or slow and can affect small
or large areas; (2) climate and hydrography play an
overwhelmingly important role in determining the
major outlines of marine communities, with biological
interactions specifying the details of community
structure within the constraints lowed by the physical
environment; (3) whatever regularities we may in the
structure of assemblages of marine species are within
the context c€ a very changeable environment so
generalizations and trends will be hard to find and
define.
14.2 Daily Changes: Vertical Migration
14.21 Nature and Stimuli in Vertical Migration
In sediments, motile blue-greens, dinoflagellates, and
diatoms may daily or tidally governed excursions of a
few millimeters up to the sediment surface (Gallagher
and Daiber, 1974; Eaton and Simpson, 1979).
In the water column, vertical migration may involve
dinoflagellates. zooplankton, or fish, and the vertical
movements may span tens to mazy hundreds of meters.
Some photosynthetic dinoflagellates migrate upward as
light begins to increase at dawn, and travel distances up
to 50 m. dinoflagellates travel at speeds of 1—2 x 10 -2
cm sec -I , sufficient to overcome the upwelling velocity
in the Baja California coastal
(Blasco, 1978) but perhaps not where upwelling is more
intense. Faster currents and changes in the density of
water can hinder migration dinoflagellates. Different
species of dinoflagellates have characteristics
sensitivities to light intensity; this results in a different
pattern of vertical migration and different temporal and
vertical distribution of the species
In zooplankton, vertical migration involves an active
upward movement of most species as night approaches
(Fig. 14-2) The speeds required are substantial; salps,
for instance, are most abundant near the during the
night, and may be as deep as or deeper by day;
swimming speeds to cover these distances roust be 5—
10 min ( et al., 1979).
Feeding by adult zooplankton usually takes place at
night in the shallow layers. At dawn there is a return to
the daytime depths, where individuals idle until the
next dusk (Longhurst, 1976).
In any parcel of the sea not all species of plankton
migrate to the sarrz extent or synchronously, and young
stages tend to be less active migrators. Tows taken in
the North Atlantic show different patterns of vertical
migration in adult copepods (Fig. 14-2). One species
shows a lack migration, with only a very short
downward excursion at night, A second species does
migrate downward about 350 m by day but newly
molted individuals remain near the surface. A third
species shows a well-defined vertical movement byt the
entire adult population. Thus the assemblage of
populations found at any one depth varies markedly
over a daily interval: behavioral changes on scale of a
day affect the structure of the community.
Near the surface, light is the stimulus that starts upward
migrations; after dusk there is commonly a passive
sinking as night brings darkness. At the next dawn there
may be a reversed short period of ascent before their
downward movement begins. On bright, moonlit nights
these short episodes of sinking after the migrants reach
the surface are reduced, indicating that light intensity
does govern this behavior (Longhurst, 1976). It is not
clear what cues are used for vertical migration by
zooplankton located at depths below 750— 1000 m,
since organisms supposedly cannot detect downwelling
light at such depths (Clarke and Denton, 1962) (cf. Fig.
2-3), yet 25 of 207 species collected between 600 and I ,
700 m showed vertical migration cued to day—night
cycles. Either organisms are extremely sensitive to some
aspect of light or other cues are used.
14.22 Reasons for Vertical Migration
There must be some important advantage to the
migratory behavior, since the vertical excursions
frequently take the plankton between and 50,000 times
their body lengths and since for most zooplankton
species it would seem preferable to feed continuously.
There are a number of hypotheses about why vertical
migration takes place. None is proven, but it seems
likely that migration has multiple causes.
14: The Structure of Marine Communities Over Time
Avoidance, of competition. Dinoflagellates can migrate
from surface waters to deeper layers. Dinoflagellates
have a relatively high light saturation curve (Fig. 2-5), so
they can actively photosynthesize near the surface
where other producers may be inhibited; at night they
can migrate downward to deeper waters where
nutrients are not as depleted as at the surface. Since
the abundance of other phytoplankton may be lower in
deeper water, migration may lessen competition.
Avoidance of predation. Upward migration at dusk
brings zooplankton to the surface layers during the
period when their visual predators are at a
disadvantage. Many migrants, however, move only
short distances, and spend the day at depths where
there is enough light for predators to see prey. In
addition, many migrants go deeper than might be
required just to escape predators; last, luminescent
migrants are common. On the other hands Ohman et al.
(1983) find a ' 'reverse" migration of the copepod
Pseudocalanus in a temperate fiord* presumably an
escape response to "normally" migrating predators
Pseudocalanus. The "reverse" migration is not seen
during _pcriods when predators are less abundant. This
suggests a strong influencecö predation on migratory
activity.
Horizontal transport. Vertical migration, both on a daily
and on a seasonal time scale may aid a population both
in maintaining-its location in a very dynamic
environment and in dispersal. It is often the case that
different layer of water travel in different directions.
Galatheid crabs transported by the California Current to
tropical waters have a potential way back by migrating
into an undercurrent moving in an opposite direction.
Similar situations are found in currents of the Styait of
Gibyaltar (Frassetto et 1962). in two-layered estuaries
(Bosch and Rowland Taylor, 1973), and in upwelling
(Wroblewski, 1982),
Energetic- advantages. Energetic expenditures by
animals are strongly correlated to ambient
temperatures, as we saw in Chapter 7. so that part-time
residence in deeper, colder water reduces energy
losses. Feeding is easier near the surface, where food
particles are most abundant. Vertical migration could
thus allow feeding while minimizing respiratory
expenditures. Migration is not very energetically costly
since swimming by zooplankton is not a very energy
demanding activity (Haury and Weihs, 1976). This is
especially so in some salps, where the organisms swim
actively all day long and feeding, locomotion, and
respiration all are accomplished by the same motion
(Wiebe et al. 1979), Growth is. however, slower at
lower temperatures, so the energy bonus clue to
vertical migration must be enough to compensate for
both locomotory expenditures and lower growth rate
(McLaren, 1963), The reduced respiration due to lower
temperatures (Section 7.321) may allow a reallocation
of energy into growth (Section 7.431), so that copepods
that migrate may grow to a larger size. Since larger
animals have increased fecundities (Section 4.21),
migration has a selective advantage (McLaren, 1974).
The energy bonus is. in theory at least, larger at high
temperatures so more migrant species are expected in
tropical waters where the surface water is warm is
strong contrast to deeper layers. At higher latitudes the
vertical profile temperatures are more uniform and
fewer vertical migrators can be expected (McLaren,
1963). These patterns seem to be true to some extent
but have not been proven. The argument may hold for
seasonal as well as daily vertical migrations.
Enright (1977) developed the energy-advantage idea
and suggested that nocturnal grazing by copepods
might provide a greater gain of energy than continuous
feeding, if three conditions were met: (l) if
photosynthesis is high enough so that the algal biomass
accumulates appreciably from dawn to dusk; (2) if
metabolism of copepods is lowered while they are in
the deeper layers; and (3) if the grazing rate of the
zooplankton is higher than average in returning to the
surface. From earlier discussion (Sections 1.4, 6.222,
7.321 and 5.221) we know that the three conditions are
reasonable. The relative importance of this intermittent
feeding version of the energy advantage hypothesis
versus the predator hypothesis can be tested by noting
when the zooplankton arrive at the surface, If migration
is a response 'to escape visual predators, arrival at the
surface should take place after sunset. If grazers begin
feeding in surface water a few hours before sunset,
they may gain a significant advantage; and the predator
hypothesis would seem to be less applicable. Enright
kind Honneger (1977) examined the timing of arrival of
copepods in surface waters off California. In one
sampling series carried out during spring, when
copepod density was high, they found that the onset of
migration began 1-2hr before sunset. In the summer,
when predators were less numerous migration began 1
—2 hr after sunset. Thus, at different times in the
season, different factors may affect vertical migration It
'is unlikely that there is a simple answer to the question
of why zooplanklon migrate.
Short-term phenomena such as vertical migration make
the concept of a plankton community a very diffuse
one. There are only tenuous associations among the
species present at any one depth, since some migrate
vertically and some do not, Further, there are
associations vertically up and down the water column,
since species of deep water may feed or compete with
surface species when the latter are in the deep phase of
their migration cycle. Thus, where vertical migration is
important. Biological processes at the physiological,
behavioral, and populational levels combine to make it
very difficult to find community properties.