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The Canonical Distribution of Commonness and Rarity: Part I F. W ...

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Spring 1962<br />

CANONICAL DISTRIBUTION<br />

breeding season most passerine birds are, o~vi~lg <strong>of</strong> individuals without accumulating many speto<br />

their "territorial" propensities, over-regularized<br />

spatially, <strong>and</strong> therefore on a small area they are<br />

over-rqgularized in abundance. (See below, under<br />

Thomas, Hicks, Williams, I'dalkinshaw, etc.) .<br />

cies. Thus, in order to deal at all with the<br />

But in the winter they "flock" <strong>and</strong> become<br />

"clumped," <strong>and</strong> it is not solely the rarer forms<br />

that do this.<br />

Plants are perhaps the most obviously clumped<br />

material, <strong>and</strong> sometimes approximate to pure<br />

cultures, as with Hopkins' (1955) Zostera community.<br />

However, this can be matched among<br />

colonially nesting birds, like Beebe's ( 1924) "pure<br />

culture" <strong>of</strong> boobies in the Galapagos Isl<strong>and</strong>s. or<br />

the Emperor Penguin (Apfertodytes forsteri) in<br />

Antarctica. <strong>The</strong>refore it seems to me that we<br />

are justified in examining the properties <strong>of</strong> r<strong>and</strong>omly<br />

distributed populations, <strong>and</strong> treating them<br />

as a "norm," <strong>and</strong> we nlay treat contagion, whether<br />

positive or negative, as introducing a disturbance,<br />

modification, or perturbation into our calculations.<br />

Notwithst<strong>and</strong>ing Hairston's ( 1959) comment<br />

that, so long as vie are dealing with a single community,<br />

"departure from r<strong>and</strong>omness increases<br />

with sample size," I thillk we are justified in<br />

assuming that in this paper we shall rarely be<br />

dealing .n-it11 a single community, but with what<br />

he calls heterogenous material, in which departure<br />

fro111 r<strong>and</strong>omness decreases with increasing saniple<br />

size. Thus in order to study contagion in our<br />

present context we must examine the properties<br />

<strong>of</strong> rather small samples.<br />

Since in the present paper we are concerned<br />

with abundance-distributions <strong>and</strong> want to know<br />

whether they conforn~ to the lognormal type, <strong>and</strong><br />

in particular to the canonical lognormal, a small<br />

sample for our purposes is primarily one that<br />

has rather few species, say a hundred or less. For<br />

with less than about one hundred we cannot plot<br />

the distribution graphically with much success.<br />

It is true we might use analytical methods rather<br />

than graphical ones, as being more powerful tools,<br />

but statistical fluctuations are not thereby prevented<br />

from confusing the issue, <strong>and</strong> so in Preston<br />

(1957) I suggested that an adequate sample called<br />

for something like a minimum <strong>of</strong> 200 species <strong>and</strong><br />

something like a mitlimum <strong>of</strong> 30,000 individuals.<br />

Very little in the way <strong>of</strong> plant material meets<br />

the requirement <strong>of</strong> 200 species. or eveti 100. <strong>and</strong><br />

much <strong>of</strong> it, in fact most <strong>of</strong> it, appears to have less<br />

than 50, <strong>of</strong>ten much less. <strong>The</strong> individuals may<br />

be very numerous, especially in our northern latitudes<br />

where the floras are poor compared with the<br />

tropics (Cain <strong>and</strong> Castro, 1959) but, because<br />

the majority <strong>of</strong> plant distributions are "positively"<br />

contagious, we have an abnormally high ntltnber<br />

203<br />

published literature on plant distributions, i.e.,<br />

their relative abundances in a "community" or<br />

their Species-Area curves, we have to compile<br />

a tabulation <strong>and</strong> make a graph <strong>of</strong> what to expect <strong>of</strong><br />

samples containing less than 100 species.<br />

Criteria for s~~lall sar?tples<br />

With small samples in this sense, it is easier<br />

<strong>and</strong> perhaps more accurate to reduce them to<br />

graphs, <strong>and</strong> the easiest computation to make concerns<br />

the (logarithmicj st<strong>and</strong>ard deviation o.<br />

<strong>The</strong>refore in Table I9 below we use the methods<br />

TABLEIX. Properties <strong>of</strong> small canonical ensembles<br />

I<br />

I<br />

Number <strong>of</strong> Logarithmic st<strong>and</strong>ard Ratio <strong>of</strong> individuals<br />

species S deviation a (octaves) to species I/N or<br />

more properly I/mN<br />

<strong>of</strong> our first section to get estimates <strong>of</strong> the value<br />

<strong>of</strong> 5 <strong>of</strong> 1,'mK ior canonical ensembles ("universes")<br />

with less than 100 species.<br />

<strong>The</strong>se values shorrld be regarded as approximate<br />

only. <strong>The</strong>y are graphed in Figure 17.<br />

On that same graph are shown a number <strong>of</strong> experimental<br />

points exhibiting contagion, positive <strong>and</strong><br />

negative, which will he discussed later. It should<br />

be understood that these figures relate to cornplete,<br />

non-truncated, canonical ensembles; but<br />

where small samples seem not to be too severely<br />

truncated we can compute o as if the distribution<br />

were normal in order to get a rough picture <strong>of</strong> the<br />

effects <strong>of</strong> coritagion <strong>and</strong> the "distortion" produced<br />

thereby. 11-e shall see later that the observed<br />

departure <strong>of</strong> s from its expected or canonical<br />

value gives an estimate <strong>of</strong> the degree <strong>of</strong> contagion,<br />

as indeed is otherwise obvious, <strong>and</strong> that contagious<br />

distributions do not end at the crest <strong>of</strong> the<br />

"Individual Curve."<br />

Skcrcwess us n criterion<br />

It is very likely that contagious distributions<br />

are soniewhat skewed, the mode lying to the right<br />

<strong>of</strong> the mean, i.e. at a higher abundance <strong>of</strong> individuals<br />

per species than the mean for negative<br />

contagion. This criterion, being subject to com-

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