ABD 3e Chapter 8, Section 4
By the end of this lecture, you should be able to:
If events are not independent or the rate is not constant, counts will not follow a Poisson distribution.
Instead, events may appear:
The Poisson distribution therefore represents a model of random spatial or temporal structure.
Suppose events are randomly distributed in space:
Divide the area into equal grid cells
Count events in each cell
Tabulate the frequencies of counts
If events are random, the distribution of counts per cell will follow a Poisson distribution.
Once we assume a Poisson process, we can compute exact probabilities for observing different counts:
\[ \operatorname{Pr}[X \text{ successes}] = \frac{e^{-\mu}\mu^{X}}{X!} \]
\(\mu\) is the mean number of independent successes in time or space (expressed as a count per unit time or a count per unit space, making this is a rate)
\(e\) is the base of the natural log, a constant ≈ 2.718
Assume points are distributed randomly
Divide area into a grid
Count points per square
Frequency of counts would follow a Poisson probability distribution
So far, we have described a model.
Now we use the Poisson distribution as a null hypothesis:
\(H_0\): Events occur randomly in time or space
Observed counts follow a Poisson distribution
This is a Poisson Goodness-of-Fit Test
Workflow when testing whether counts follow a Poisson distribution:
For counts to follow a Poisson distribution:
If these assumptions fail, counts may be clumped or dispersed.
Biological question:
Do extinctions occur randomly through geological time, or are there intervals with unusually high extinction rates?
If extinctions are random, counts per time interval should follow a Poisson distribution.
These are the observed counts we will compare to Poisson expectations.
Raup DM, Sepkoski JT. 1982. Mass extinctions in the marine fossil record. Science 215: 1501-1503.
If extinction events are random in time:
Therefore:
This is a goodness-of-fit test.
Step 1: Estimate the Poisson rate parameter ( \(\mu\) ) under \(H_0\).
We estimate \(\mu\) using the sample mean number of extinctions per interval.
| Number of extinctions | Frequency |
|---|---|
| 0 | 0 |
| 1 | 13 |
| 2 | 15 |
| 3 | 16 |
| 4 | 7 |
| 5 | 10 |
| 6 | 4 |
| etc. | … |
\[ \bar{X}=\frac{(0\times0)+(1\times13)+(2\times15)+\dots}{76} \\ = 4.21 \]
We can use this expected mean \(X\) in place of \(\mu\) in the formula for the Poisson distribution to generate the expected frequencies
\[ \operatorname{Pr}[X \text{ successes}] = \frac{e^{-\mu}\mu^{X}}{X!} \]
\[ \operatorname{Pr}[3 \text{ extinctions}] = \frac{e^{-4.21} \cdot 4.21^{3}}{3!} \]
\[ \operatorname{Pr}[3 \text{ extinctions}] = 0.1846 \]
\[ \operatorname{E}[3 \text{ extinctions}] = 76 \times 0.1846 = 14.03 \]
We subtract 1 degree of freedom because:
Probabilities must sum to 1
We estimated one parameter (μ) from the data
This reduces the available independent information.
\[ df=(\operatorname{number of categories})-1-(\operatorname{number of parameters}) \\ df = 8 - 1 - 1 = 6 \]
Critical value for chi-square with 𝑑𝑓=6 at 𝛼=0.05 : 𝟏𝟐.𝟓𝟗
Our chi-square test statistic: 𝟐𝟑.𝟗𝟑
Do we reject \(H_0\) or not?
Test statistic: 23.93
Critical value (df = 6, α = 0.05): 12.59
Because 23.93 > 12.59:
Statistical conclusion:
Biological interpretation:
Extinctions are not occurring at a constant random rate.
There are intervals with unusually high extinction rates.
This pattern is consistent with episodic mass extinction events.
An alternative diagnostic for randomness:
Compare the variance to the mean.
Variance ≈ Mean → Consistent with Poisson
Variance > Mean → Clumped pattern
Variance < Mean → Dispersed pattern
This provides a quick check before formal testing.
Binomial distribution:
Poisson distribution:

BIOL 275 Biostatistics | Spring 2026