Chapter 9. Relative Abundance and Diversity
In addition to the high diversity of the coral reef ecosystem, a feature of this ecosystem is the large number of rare species within each taxonomic group. The general relation between the number of species and the number of individuals in a sample of a population was discussed by Fisher, Corbet and Williams (1943), who commented that species are not equally abundant, even under conditions of considerable uniformity. They went on to state that the majority of species are comparatively rare while only a few are common.
It is not known whether the rarity of a species is indicative of its low competitive ability or alternately whether the species is restricted to specialised microhabitats with excess recruitment eliminated by predators (Hairston, 1959; Kunin and Gaston, 1993). The relative abundances of the species in a diverse assemblage are often distributed over many orders of magnitude. As a result, qualitative representations of abundance such as common, moderately abundant or rare must be arbitrary in their assignment.
Many different mathematical models have been proposed to describe satisfactorily the relationship that exists between the relative abundances of different species in an assemblage. While each model has been criticised extensively (Hurlbert, 1971; Abbott, 1983; Connor and McCoy, 1979; Connor, McCoy and Cosby, 1983; Martin, 1981; McGuiness, 1984; Pielou, 1981; Sughihara, 1981), each attempts to quantify the degree of variation in the relative abundances of the different species. The most noticeable result of this abundance variation is the different rates at which species accumulate with increased sampling in different assemblages.
The population density of each species and the relation between sample area and the number of individuals in the sample was calculated in Chapter 4. The relation between sample area and the total number of species in the sample (the species : area curve) was also calculated from the traverse data. The cumulative number of species was compared with the cumulative area of the traverses (starting at the completion of Traverse 1 and continuing through to the completion of Traverse 72). This comparison was also undertaken with the natural logarithm of the cumulative area of the traverses.
Shannon’s Evenness Index (see Pielou, 1981) which is the expression (S P(log P)) / log S, where P is the proportion of each species in the community, and S is the total number of different species, is often used to display the relative richness of various communities. Shannon’s Evenness was calculated for each traverse individually and cumulatively starting with Traverse 1 and ending with Traverse 72.
The relation between the numerical abundance of each species and the rank abundance of each species was calculated by ordering the numerical abundance from most common (rank 1) to least common (equal rank 20 for five species). Percent relative abundance was the ratio of the numerical abundance of each species to the total asteroid abundance.
Table 9.1 lists the numerical, relative and rank abundances of each species located on the intertidal traverses. Figure 9.1a graphs the relation between the numerical abundance of a species and its rank abundance. Figure 9.1b graphs the relation between (log) relative abundance and rank abundance. Figures 9.2a,b graph the species : area and species : (log) area relation. Figures 9.3a,b graph the relation between Shannon’s Evenness and cumulative area and cumulative (log) area. Natural logarithms were used in all these calculations. Shannon’s Evenness as a measure of diversity has the advantage that the index is a ratio of attained diversity over maximum possible diversity and is therefore independent of the base of logarithm which has been chosen.
The numerical abundance, relative abundance and abundance rank of inter-tidal asteroids at Heron Reef.
SPECIES NUMERICAL RELATIVE RANK Culcita novaeguineae 15 ** 13 Asteropsis carinifera 3 * 19 Dactylosaster cylindricus 1 * 20 Fromia elegans 16 ** 12 Fromia milleporella 1 * 20 Gomophia egyptiaca 6 * 16 Linckia guildingii 116 *** 8 Linckia laevigata 509 *** 3 Linckia multifora 522 *** 2 Nardoa novaecaledoniae 326 *** 5 Nardoa pauciforis 187 *** 7 Nardoa rosea 1 * 20 Ophidiaster armatus 4 * 17 Ophidiaster confertus 4 * 17 Ophidiaster granifer 116 *** 9 Ophidiaster lioderma 1 * 20 Ophidiaster robillardi 24 ** 10 Asterina anomala 17 ** 11 Asterina burtoni 208 *** 6 Disasterina abnormalis 500 *** 4 Disasterina leptalacantha 7 * 14 Tegulaster emburyi 1 * 20 Echinaster luzonicus 1402 **** 1 Coscinasterias calamaria 7 * 14 * Very rare <10 :** rare 11-100 :*** common 101-1000 :**** abundant>1000
The generally low abundances of most of the species of starfish at Heron Reef precluded the use of quadrats in general sampling. Because the traverse method will miss many cryptic individuals and provide only an approximate area measurement, the species diversity and species accumulation figures are only approximate. It would appear from Table 4.1 that most species occurred at a density that was less than one individual per hectare, with many species being far less abundant. It should be noted that traverse sampling will underestimate the density of all cryptic species, and will also fail to detect species that are both rare and cryptic.
McGuiness (1984) suggested that the use of species : (log) area or (log) species : (log) area for the display of the species : area relationship should be based on the underlying relative abundances of the species. The slope of the species : (log) area relationship, the slope of the (log) relative abundance : rank abundance relationship and Shannon’s Evenness index are all indices of diversity. These allow a direct comparison to be made between different assemblages. Not only do these indices consider the number of species, they also express the inherent range of abundance between most common and least common within the assemblage (Connor and McCoy, 1979; Connor and Simberloff, 1979; Connor et al., 1983).
Figures 9.1a,b show that the four most common species account for 70% of the total number of individuals in this assemblage. However, even Echinaster luzonicus, the most abundant species, had an average density of only 16 specimens per hectare. Of the 24 species of asteroid that occurred in the traverse samples, five species occurred only once. Presumably the species which were not found during this study, but which are known from the locality, occur with even less frequency than these five. Less than ten specimens of each of another six species were located on the intertidal traverses. Hence, 11 of the 24 species are regarded as very rare. Less than 25 specimens of another three species were found and these are regarded as rare. Thus a majority of the asteroid species found at Heron Reef are rare or very rare.
The slope of the regression line in Figure 9.1b is a measure of the diversity of this asteroid assemblage. The steeper the line the greater the range of relative abundance within a certain group of species. The less equal the relative abundances, the lower the diversity as measured by most diversity indices. Community studies often show a log-normal relationship in relative abundance, in which most species occur with close to the average abundance (Pielou, 1981). This assemblage of coral-reef asteroids does not clearly demonstrate this relationship, but this result may be attributable to an inadequate number of both species and individuals in the present study. The order of the species in Figures 9.1a,b is that of numerical abundance. If biomass or some other parameter was chosen as a measure of abundance, then the order of the species may change but the slope of the regression line might not alter greatly.
Figures 9.2a,b illustrate the species : area curve for the Heron Reef asteroid assemblage. The slope of the (log) area regression line is independent of the units used to measure area. Whether they be square metres or hectares, providing the habitat continues, the species will accumulate at a rate determined only by the relative abundances of the species in the assemblage. If there is some finite species pool which obviously cannot be exceeded, then the curve will become asymptotic.
The pronounced dips in Figures 9.3a,b are a result of small scale patchiness in the distribution of Echinaster luzonicus and Disasterina abnormalis. After continued sampling, the effect of this high localised abundance was rendered insignificant in the total diversity.
Figures 9.1a to 9.3b all relate to the one ecological parameter, namely the relative abundances of the species within this assemblage. This will determine the rate at which the species accumulate in a species : area curve, as well as the diversity as measured by most diversity indices.
The richness of the coral-reef asteroid assemblage at Heron Reef is unable to be compared directly with that of other coral-reef asteroid assemblages either on the Great Barrier Reef or elsewhere. This is because the extent of sampling has not been quantified in the majority of biogeographical studies. Because the area sampled determines the number of species in a sample of any assemblage (Fisher, Corbet and Williams (1943), the large number of species found at Heron Reef may be a result of the intensive sampling. Even so, it would appear from the linearity of Figures 9.2b that additional species of starfish occur intertidally at Heron Reef, but these species are either extremely rare or cryptic.
It is apparent that Heron Reef carries a rich and diverse asteroid fauna, 24 species belonging to six families having been found intertidally in 120 hectares during this study. The linearity of the species : (log) area relationship for the intertidal asteroid assemblage at Heron Reef indicates that additional species are still to be found. Indeed, Mithrodia clavigera was located subsequent to the traverses and Endean (1956) found three species (Acanthaster planci, Ophidiaster watsoni and Anseropoda rosacea) in the area of the traverses that were not found during the current study.