4.7.

2 Wall structure
Marsh and mangal assemblages plot throughout the ternary diagram (Figure
4.18). Many have 100% agglutinated walls, especially those from high marshes and
from marshes furthest from the open sea. Hyaline forms appear on low marshes and
marshes closer to the open sea. Brackish marshes plot mainly along the agglutinated–
hyaline side and hypersaline marshes along the hyaline–porcelaneous side. Marshes
that vary seasonally from brackish to hypersaline (e.g., Texas) plot along both these
sides. Brackish estuaries and lagoons plot mainly along the agglutinated hyaline side
with only a small porcelaneous component (Figure 4.19). Normal marine and
hypersaline examples plot along the hyaline–porcelaneous side with varying amounts
of agglutinated walls. The same applies for deltas (Figure 4.20). In fjords, the normal
marine deep-water assemblages plot along the agglutinated–hyaline side while the
variable surface waters often have some porcelaneous forms.

4.7.3 Standing crop in 0–1 cm sediment layer

Samples with a count of fewer than 50 have been excluded from con-
sideration so it is not possible to calculate average values. There are some zero
values in all environments. Two features characterise the data: extreme varia-bility
3
within a small area; maximum values often several thousand individuals 10 cm . For
marshes (Table 4.1), with the exception of the Mediterranean (maximum 330
3 3
individuals 10 cm ), maximum values exceed 1000 individuals 10 cm throughout
3
the world. The typical values are several hundred 10 cm . For estuaries and lagoons,
3
maximum values in Europe are 2000–3000 indivi-duals 10 cm . However, values
are typically low in the Arctic, Exe estuary and Venice (<50) but >200 in the Hamble
and northern Spain. In Atlantic North America, maximum values are 260–756 with
3
typical values of 50–200 indivi-duals 10 cm . Pacific lagoons have maximum 956
3
and typically 100–200 indi-viduals 10 cm . There is a marked contrast between the
Ebro delta (maximum 240, typically 10–100) and the Mississippi delta (maximum
3
8240, typically 80– 500 individuals 10 cm ) reflecting the nutrient differences
between the two areas. Fjords in Europe are very variable according to degree of
oxygenation. Maximum values sometimes exceed 1000 but may be as low as 100
3
and typical values may be <50 or range up to several hundred individuals 10 cm .
Marginal marine environments

Figure 4.19. Ternary plots of walls for estuaries and lagoons (triangle corners represent 100% of the labelled
component: A = agglutinated, P = porcelaneous, H = hyaline).
4.8 Summary

Although ecologists distinguish between estuaries, lagoons and deltas on

geomorphological and sedimentological criteria, it is apparent that these distinctions are
not important for the fauna. In general species diversity is low in extreme environments
(brackish, hypersaline marshes, lagoons and estu-aries) but ranges a little higher in
normal marine lagoons. In terms of wall structure, whereas in marshes there are some
assemblages that comprise mixtures of agglutinated and porcelaneous tests (with few
hyaline forms), that is not the case for lagoons, estuaries, deltas or fjords. The general
pattern is that brackish subtidal environments have assemblages with a mixture of
agglutinated and hyaline walls. Hypersaline environments (found only in warm climates)
have mixtures of porcelaneous and hyaline walls.
114 Marginal marine environments

3
Table 4.1. Summary of standing crop data for the 0–1 cm sediment interval (values individuals 10 cm ).

Environment Maximum value Typical value Location

Marsh 1798 200–1000 UK

1252 100–200 Spain
330 50–100 Mediterranean
2464 50–1000 Atlantic USA
2450 100–200 Gulf of Mexico
1652 >100 Pacific North America
10908 300–5000 Japan
Estuaries 3404 >200 Europe
756 50–200 Atlantic USA
2784 50–400 Gulf of Mexico
376 10–75 South America
1652 50–300 Pacific North America
Deltas 240 10–100 Spain
8240 80–500 Gulf of Mexico
Fjords 2109 207–500 Europe

Environments that change seasonally from brackish to hypersaline have assemblages that
accordingly plot along both these sides of the ternary dia-gram.

As can be seen from the foregoing account, major species distributions and
biogeographic boundaries are controlled primarily by a combination of tem-perature and
salinity. Species may be confined to a certain range of salinities but there are no species
found only in a single geomorphological setting such as a lagoon, estuary, delta or fjord.
However, there are certain species found only in association with vegetated marshes and
only rarely on adjacent mud-flats (Balticammina pseudomacrescens, Jadammina
macrescens, Tiphotrocha compri-mata, Trochammina inflata). Local species distributions
are determined by a plexus of abiotic and biotic controls. In those studies where authors
have used multivariate and statistical measures to determine the relationships between
abiotic variables and species there is always a great deal of co-variance and
interdependence between variables. Consequently, it is difficult to know exactly which
controls are most important. It seems likely that the importance of each variable changes
with time as it comes close to the threshold for any given species. Therefore, different
variables are significant at different times. Also some interdependent variables may
reinforce one another in their influ-ence on the fauna. Thus, salinity and temperature may
interact so the response
 4.8 Summary 115

of an organism to a change in salinity may be different at different tempera-tures, as these affect osmosis,
concentration of dissolved oxygen, density and viscocity. There are differences between non-tidal and tidal
settings. The for-mer have a much more stable temperature and salinity structure on timescales ranging from
days to weeks. Tidal areas are affected by currents and mixing so that changes in environmental parameters are
continuous even if diurnally cyclic. Therefore some fjords and the Baltic have certain species adapted to
constant conditions (Ammotium cassis living at the halocline). The standing crop is controlled by the
availability of suitable food, dissolved oxygen (if dysoxic) and the degree of predation. Although maximum
3
values sometimes reach several thousand individuals 10 cm typical values are generally 50 to a few hundred.
Several studies point to selectivity for particular types of food. In general, there is non-competitive feeding
with each species taking a different part of the potential food supply.

In conclusion, it can be seen that although the general controls limiting marginal marine foraminiferal assemblages
have been established there is still much to be learned about the operation of factors at a local level. Future progress in
unravelling this will come from a combination of carefully con-trolled laboratory experiments to understand the
physiological response to a forcing factor and field studies targeting specific themes