Stage-structured population growth

The exponential growth model: assumptions

  • No immigration or emigration (Closed population)

  • Constant birth and death rate (\(b\) and \(d\) don’t vary with \(N\))

  • No variation within population (all individuals have similar \(b\) and \(d\))

    • This implies that \(b\) and \(d\) don’t vary with age or stage
  • Continuous population growth without time lags
    (e.g. no seasonality)

Organisms with stage structure

Photo by Mike Erskine on Unsplash

Modeling structured populations

Consider a bird species where each (female) adult lays \(10\text{ eggs}\). Only \(10\%\) of eggs successfully hatch into juveniles, and \(50\%\) of juveniles survive and mature into adults. Adults die after reproduction.

  • How can we represent this in a diagram?

Photo by Mike Erskine on Unsplash

Consider a short-lived plant that lives for 6 years. Each year, only \(1\%\) of seeds survive and germinate into seedlings. (The other seeds are eaten by squirrels and jays). \(50\%\) of seedlings survive to a juvenile stage, and \(95\%\) of juveniles transition into reproductive sub-adults. Each sub-adult makes \(10\text{ seeds}\) per year, and \(95\%\) of sub-adults transition into fully-mature adults. Each mature adult makes \(450\text{ seeds}\) per year, and \(10\%\) of adults survive to a post-reproductive adult stage, which are incapable of making new seeds.

  • Draw the transition diagram.

Diagrams are helpful, but numbers are better

Transition matrices

These transition diagrams can be represented mathematically as transition matrices.

For a population with \(n\) stages, we can capture the demographic transitions in an \(n\mathrm{-by-}n\) matrix.

  • Each element captures the contribution from the current column, to the current row.
    • The top row always reflects new births
    • The first (leftmost) column reflects the youngest individuals
    • The last (rightmost) column reflects the oldest individuals

Worked example

Consider a bird species where each (female) adult lays \(10\text{ eggs}\). Only \(10\%\) of eggs successfully hatch into juveniles, and \(50\%\) of juveniles survive and mature into adults. Adults die after reproduction.

\[ \begin{bmatrix} ~ & ~ & ~ & ~ & ~& ~ & ~ \\ ~ & ~ & ~ & ~ & ~& ~ & ~ \\ ~ & ~ & ~ & ~ & ~& ~ & ~ \\ ~ & ~ & ~ & ~ & ~& ~ & ~ \\ ~ & ~ & ~ & ~ & ~& ~ & ~ \\ ~ & ~ & ~ & ~ & ~& ~ & ~ \end{bmatrix} \]

Worked example

Consider a bird species where each (female) adult lays \(10\text{ eggs}\). Only \(10\%\) of eggs successfully hatch into juveniles, and \(50\%\) of juveniles survive and mature into adults. Adults die after reproduction.

\[ \begin{bmatrix} 0 & 0 & 10 \\ 0.1 & 0 & 0 \\ 0 & 0.5 & 0 \end{bmatrix} \]

Your turn

  • Consider a species of fish whose individuals live for 3 years (age classes 0, 1, 2, and 3). Newborn fish have a \(30\%\) survival rate to year 1; year 1 fish have a \(80\%\) survival rate to year 2; year 2 fish have a \(50\%\) survival rate to year 3; and all fish die in their third year.

  • Newborn fish are sexually immature and cannot give birth. Year 1 fish can give birth to \(1\) newborn per year; Year 2 fish can give birth to \(8\) newborns per year; and Year 3 fish can only give birth to \(0.5\) newborns per year.

  • Draw a transition diagram and write the transition matrix for this population.

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \]

The value of a transition matrix

  • The product of the current population distribution and the transition matrix tells you the future population distribution

The value of a transition matrix

  • From our worked example, recall the transition matrix

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \]

  • At time \(t=0\), the population has \(52\) individuals in stage 0, \(10\) in stage 1, \(15\) in stage 2, and \(30\) in stage 3.

  • What is the expected distribution of individuals at \(t=1\)?

The value of a transition matrix

Matrix product of the transition matrix and the current distribution:

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \times \begin{bmatrix} 52 \\ 10 \\ 16 \\ 30 \end{bmatrix} \]

The value of a transition matrix

Matrix product of the transition matrix and the current distribution:

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \times \begin{bmatrix} 52 \\ 10 \\ 16 \\ 30 \end{bmatrix} = \begin{bmatrix} 153 \\ 15 \\ 8 \\ 8 \end{bmatrix} \]

The value of a transition matrix

For timestep \(2\):

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \times \begin{bmatrix} 153 \\ 15 \\ 8 \\ 8 \end{bmatrix} \]

The value of a transition matrix

For timestep \(2\):

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \times \begin{bmatrix} 153 \\ 15 \\ 8 \\ 8 \end{bmatrix} = \begin{bmatrix} 83 \\ 45.9 \\ 12 \\ 4 \end{bmatrix} \]

The value of a transition matrix

For timestep \(3\):

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \times \begin{bmatrix} 83 \\ 45.9 \\ 12 \\ 4 \end{bmatrix} = \dots \]

The value of a transition matrix

  • We can keep iterating over and over (and over), or….
  • Calculate the \(\mathrm{eigenvalue}\) of the transition matrix
  • The dominant eigenvalue reflects the long-term growth rate (\(\lambda\)).

\[ \begin{bmatrix} 0 & 1 & 8 & 0.5 \\ 0.3 & 0 & 0 & 0 \\ 0 & 0.8 & 0 & 0 \\ 0 & 0 & 0.5 & 0 \end{bmatrix} \xrightarrow[]{\text{Eigenvalue}} 1.33 \]

Computing eigenvalues in R

The value of a transition matrix

  • The dominant Eigenvalue represents the expected long-term annual growth rate (\(\lambda\))
    • Over the long term, \(\frac{N_{t+1}}{N_t} = \lambda\)
  • Populations grow when \(\lambda \gt 1\)
  • Populations shrink when \(\lambda \lt 1\)
  • Population size is stable when \(\lambda = 1\)

Reminders for next week

  • Read Crouse et al. 1987 paper on using transition matrices to study turtle population dynamics
  • Listen to Sea Change podcast about sea turtle population dynamics in the 2020s
  • In-class activity next week.