Biodiversity and Conservation 13: 9–52, 2004.
© 2004 Kluwer Academic Publishers. Printed in the Netherlands.
The role of density regulation in extinction
processes and population viability analysis
KLAUS HENLE
1,∗
, STEPHEN SARRE
2
and KERSTIN WIEGAND
3,4
1
Department of Conservation Biology and Natural Resources, Centre for Environmental
Research, UFZ Leipzig-Halle, Permoserstr. 15, D-04318 Leipzig, Germany;
2
Applied
Ecology Research Group, University of Canberra, Canberra, ACT 2601, Australia;
3
Department of Ecological Modelling, Centre for Environmental Research, UFZ
Leipzig-Halle, Permoserstr. 15, D-04318 Leipzig, Germany;
4
Present address:
Institute of Ecology, Jena University, Dornberger Str. 159, D-07743 Jena, Germany;
∗
Author for correspondence (e-mail: klaus.henle@ufz.de; fax: +49-341-2353191)
Received 29 June 2002; accepted in revised form 15 March 2003
Abstract. We review the role of density dependence in the stochastic extinction of populations
and the role density dependence has played in population viability analysis (PVA) case studies.
In total, 32 approaches have been used to model density regulation in theoretical or applied
extinction models, 29 of them are mathematical functions of density dependence, and one
approach uses empirical relationships between density and survival, reproduction, or growth
rates. In addition, quasi-extinction levels are sometimes applied as a substitute for density
dependence at low population size. Density dependence further has been modelled via explicit
individual spacing behaviour and/or dispersal. We briefly summarise the features of density
dependence available in standard PVA software, provide summary statistics about the use of
density dependence in PVA case studies, and discuss the effects of density dependence on
extinction probability. The introduction of an upper limit for population size has the effect
that the probability of ultimate extinction becomes 1. Mean time to extinction increases with
carrying capacity if populations start at high density, but carrying capacity often does not have
any effect if populations start at low numbers. In contrast, the Allee effect is usually strong
when populations start at low densities but has only a limited influence on persistence when
populations start at high numbers. Contrary to previous opinions, other forms of density de-
pendence may lead to increased or decreased persistence, depending on the type and strength
of density dependence, the degree of environmental variability, and the growth rate. Further-
more, effects may be reversed for different quasi-extinction levels, making the use of arbitrary
quasi-extinction levels problematic. Few systematic comparisons of the effects on persistence
between different models of density dependence are available. These effects can be strikingly
different among models. Our understanding of the effects of density dependence on extinction
of metapopulations is rudimentary, but even opposite effects of density dependence can occur
when metapopulations and single populations are contrasted. We argue that spatially explicit
models hold particular promise for analysing the effects of density dependence on population
viability provided a good knowledge of the biology of the species under consideration exists.
Since the results of PVAs may critically depend on the way density dependence is modelled,
combined efforts to advance statistical methods, field sampling, and modelling are urgently
needed to elucidate the relationships between density, vital rates, and extinction probability.