position may exceed the requirements of several barnacles. In such cases, space can be seen as a resource in limited supply. Another type of interference competition occurs when, for instance, two red deer stags fight for access to a harem of hinds. Either stag, alone, could readily mate with all the hinds, but they cannot both do so since matings are limited to the ‘owner’ of the harem. Thus, with exploitation, the intensity of competition is closely linked to the level of resource present and the level required, but with interference, intensity may be high even when the level of the real resource is not limiting.
In practice, many examples of competition probably include elements of both exploitation and interference. For instance, adult cave beetles, Neapheanops tellkampfi, in Great Onyx Cave, Kentucky, USA compete amongst themselves but with no other species and have only one type of food – cricket eggs, which they obtain by digging holes in the sandy floor of the cave. On the one hand, they suffer indirectly from exploitation: beetles reduce the density of their resource (cricket eggs) and then have markedly lower fecundity when food availability is low (Figure 5.2a). But they also suffer directly from interference: at higher beetle densities they fight more, forage less, dig fewer and shallower holes and eat far fewer eggs than could be accounted for by food depletion alone (Figure 5.2b).
Figure 5.2 Competition may combine elements of both exploitation and interference. Intraspecific competition amongst cave beetles (Neapheanops tellkampfi). (a) Exploitation. Beetle fecundity is significantly correlated (r = 0.86) with cricket fecundity (itself a good measure of the availability of cricket eggs – the beetles’ food). The beetles themselves reduce the density of cricket eggs. (b) Interference. As beetle density in experimental arenas with 10 cricket eggs increased from one to two to four, individual beetles dug fewer and shallower holes in search of their food, and ultimately ate much less (P < 0.001 in each case), in spite of the fact that 10 cricket eggs was sufficient to satiate them all. Means and standard deviations are given in each case.
Source: After Griffith & Poulson (1993).
5.2 Intraspecific competition, and density‐dependent mortality, fecundity and growth
5.2.1 Density‐dependent mortality and fecundity
under‐ and overcompensating density dependence
The likely effect of intraspecific competition on any individual is greater the more competitors there are. The effects of intraspecific competition are thus said to be density dependent. We see this in more detail in Figure 5.3a, which shows the pattern of mortality in the barnacle, Semibalanus balanoides, on a rocky shore in north Wales, UK, between their recruitment as settling larvae and their survival as established adults two years later. The same data have been expressed in two ways. From the upper panel it is clear that initially, as the number of recruits increased, this led to a corresponding increase in the number of surviving adults. But the number of survivors peaked at a recruit density of around 30 per cm2 and thereafter declined. We can therefore say that at densities beyond this peak, the mortality rate showed overcompensating density dependence in that increases in initial numbers led to decreases in final numbers. By contrast, before the peak, the density dependence was undercompensating in that final numbers continued to rise as the number of recruits increased.
Figure 5.3 Density‐dependent mortality. (a) Upper panel: the density of surviving adults of the barnacle, Semibalanus balanoides, in the UK as a function of the density of recruits two years earlier. Lower panel: the same data expressed as the relationship between the daily mortality rate and the density of recruits. (b) The density of surviving seedlings recruited to a population of the yellow star thistle, Centaurea solstitialis, in California as a function of the number of seeds present in the previous year.
Source: (a) After Jenkins et al. (2008). (b) After Swope & Parker (2010).
The lower panel plots the mortality rate against the initial number of recruits (mortality rate being calculated as –ln(S/R), where S and R are the number of survivors and recruits, respectively, divided by 730 (two years) to give a daily rate – similar to the intrinsic rate of natural increase, r, calculated in Section 4.7.1). We can see that at the very lowest abundances of recruits, the relationship was flat. That is, the mortality rate stayed the same and was thus density independent. There was no evidence of intraspecific competition when initial abundances were low. As those abundances increased, the slope of the relationship became positive – there was density dependence and thus evidence of competition – and that slope became steeper as the density dependence moved from under‐ to overcompensation.
exactly compensating density dependence
A similar relationship is shown in Figure 5.3b, but this time for a plant, the yellow star thistle, Centaurea solstitialis, in California, USA, relating the density of seedlings to the initial number of seeds in the soil. This time, though, at the highest seed densities, the number of surviving seedlings levelled off. The density dependence was exactly compensating: as initial density increased the mortality rate rose to counteract it.
intraspecific competition and fecundity
The patterns of density‐dependent fecundity that result from intraspecific competition are, in a sense, a mirror‐image of those for mortality (Figure 5.4). Here, though, the per capita birth rate falls as intraspecific competition intensifies. At low enough densities, the birth rate may be density independent (Figure 5.4a, lower densities). But as density increases, and the effects of intraspecific competition become apparent, birth rate initially shows undercompensating density dependence (Figure 5.4a, higher densities), and may then show exactly compensating density dependence (Figure 5.4b, throughout; Figure 5.4c, lower densities) or overcompensating density dependence (Figure 5.4c, higher
