Recognited, each corresponding to a dfilerent section of the baseline of the gure.

The section a-b

includes the species-poor vegetation found in environ ments subected to extreme conditions of stress
andor disturbance. The cor nidor containing vogctation types with the potential for relatively high
species density occupies the intermediate part of the range in biomass (b-e), whilst the ection ed
corresponds to vegetation in which species densities are suppressed by dominance Two processes can
be identified which determine the species densities attained in particular corridor environments. The
first of these has been examined in some detail earlier in this chapter and consists of the degree of
spatial and temporal variation and the resulting opportunities for complemen tary forms of exploitation
and regeneration within the environment. The sccond process, which will be considered under the next
heading, is the availabiity and rate of ingress of potential constituent species from the surrounding
landscape.

In the lower part of Figure 58 an attempt has been made to illustrate the relationship of the model to
patterns of variation in species density which can bei observed in the field. Horizontal lines have been
drawn to indicate those parts of the model which correspond to the range of conditions in particular
habitats or vexetation types It must be emphasized that although the model may be used to interpret
variation in species density within either herb or tree layers, there will be differences of detail in the way
in which the mechanisms controlling species density operate on herbs and trees. In particular, it is clear
that in thei tree layer the thresholds (a,b,c) at which various phenomena come into play along the
baseline of Figure 58 will correspond to very much higher values in biomass. Moreover, for the reasons
discussed on page 163, we may expect that there will be quantitative differences between tropical and
temperate vegetation types.

In passing, it is interesting to note that studies of species density and com munity structure in intertidal
algal communities (Dayton, 1970; Paine, 1969, 1974) and benthic algae (Lewis, 1914; MacFarlane and
Bell, 1933; Hehre and Mathieson, 1970; Sears and Wilce, 1975; Dayton and Hessler, 1972) have
recognized many of the phenomena included in the general model presented here. There is also
evidence which suggests that many of the same principles may be applied to the control of species
density in colonial animals such as corals (Branham et al. 1971; Dana et afl, 1972; Goreau et al, 1972),
shell-fish (Connell, 1961, 1972), and even some noncolonial organisms such as coral reef fish (Sale,
1977).

Reservoir effects' upon species density

So far it has been convenient to analyse the control of species density mainly in terms of the structure of
vegetation and the interactions of the component plants both with each other and with various features
of the environment. Already, however (page 16), it has been necessary to reler to the role ol an
additional factor which may exercise a major limitation on species density. This is the reservoir of
suitable species in each geographical area. Particulary in extensively-disturbed landscapes such as that of
Lowland Britain, certain habitats such as woodland or unproductive calcareous pasture or marshland
have been reduced by agricultural development to small isolated fragment. Where these fragments are
of recent origin it is frequently observed that their potential for high species density remains unfuliled
and we may suppose that this is, in part, because of the slow rate of ingress of suitable plants from the
adjacent countryside. In these circumstances, low species densities appear to be determined by the low
dispersal efliciencies of the seeds of many plants and also by the depauperate state of the surrounding
flora.

A rather different reservoir effect appears to be involved in the consistent differences in species density
which are observed in temperate regions between adjacent and structurally similar vegetation types on
calcareous and acidic soils (e.g. Plate 34). From data such as those illustrated in Figure 44, for example, it
is apparent that in Britain, regardless of habitat, species denaities in herbaceous vegetation are
consistently low where the surface soil pH is less than 40. A more specific example of the reduction in
species density associated with increasing soil acidity is presented in Figure 59 which is based upon
survey data from an area of Northern England in which there is an intimate mosaic in which acidic and
calcareous grasslands experience comparable effects of climate and and-use. From this figure it is
evident that whereas high species densities are attained over the pH range 4.0-8.0, there is an abrupt
decline on more acidic soils. When variation in species density across the soil pH range is compared with
the total number of species recorded from all the samples examined in each soil pf category it is
apparent that the fall in species density in individual samples is correlated with a progressive reduction
in the size of the species rescrvoir associated with the transition from calcicoles to calcifuges. Reference
to the Flora of the British Iules (Clapham, Tutin, and Warburg, 1962) and the Ailar of the Britih Florn
(Perring and Walters, 1962) confirms that the greater abundance of calcicoles over calcifuges in Britain
applies not only to grassland herbs but to other ecological groups such as trees, shrubs, and ruderal
plants. This suggests that the maximum species densities which can be attained in any particular
vegetation type will show a predictable relationship to soil pH. In Figure 60 an attempt has been made to
summarize the relationships between maximum speces density, maximum standing crop + litter and soil
pH in herbaccous vegetation of the British Isles.

We may conclude, therefore, that at latitudes such as that of the British 1sles, the higher species
densities supported by calcareous soils are related, in part, to the fact that in such regions calcicolous
vegetation draws upon a reservoir of species which is considerably larger than that of the calcifuges.
"The most likely cxplanation for the greater abundance of calcicoles is that these plants have evolved
mainly at lower latitudes where the etiect of low precipitation : evap oration ratios is to maintain a high
base-status in the soils. This is to suggest that the evolution of calcicoly has occurred in semi-arid
environments where, as suggested by Margalef (1968), Stebbins (1952, 1972), Stebbins and Maior
(1965), and Bartholomew, Eaton, and Raven (1973), the effects of climatic fluctuation are to maintain a
greater degree of vegetation disturbance and cnvironmental heterogeneity, with higher rates of turnover
in plant populations, all of which are conducive to relatively rapid rates of speciation and diversification
of floras.

Latitudinal gradients in the size of the reservoir of angiosperms extend from the polar reions to the
equator (Figure 61), and this fact has important implications for the control of species density in a variety
of vegetation types. In lorests, for example, it is well known that whereas in temperate regions the mean
density of trees is less than 10 species/Ha, values exceeding 100 species Ha are frequently encountered
in tropical rainforest (Longman and Janik, i 974). It scems reasonable to suggest, therefore, that the
higher densities of tree species in tropical forests are related to the larger reservoir of trees present at
low latitudes and this, in turn, may be attributed to the long and uninterrupted period over which forest
speciation and co-adaptation (Dobzhansky, 1950: an hae heen ahle to take place in equatorial habitats.

Soil pH cotegories

Figure 59 Mean (6), maximum (e), and minimum (o) species density and total number of species
represented (e) in categories of surface soil pH encountered in 593 one square metre quadrats located at
random in unmanaged grasslands distributed within an area of 2400 km in Northern England. The values
inserted on the topmost curve refer to the number of quadrats falling in each half-unit soil pH category
(Grime, 1973c). (Reproduced with permission from Ennir. Man.. I, 151-167, Copyright 1973 by Academic
Press Inc. (London) Ltd.)

Figure 60 Diagram summarizing the relationship ween surface sol pH, seasonal maximum in standing
crop+ litter, and maximmum potential species herbaceous vegetation of the British Isles