Update

Trends in Ecology and Evolution December 2011, Vol. 26, No. 12

Letters

Minimum viable population limitations ignore

evolutionary history
Stephen T. Garnett and Kerstin K. Zander
Research Institute for the Environment and Livelihoods, Charles Darwin University, Casuarina, NT 0909, Australia

Flather et al. [1] argue convincingly that calls to incorporate a median minimum viable population (MVP) size of
5000 individuals into considerations of conservation funding [2] should be ignored. This is especially important
given the apparent biases within the data set used to
develop the median MVP on which the policy advice
was based [3]. First, the data set is biased towards publications in which MVP is explicit. Selection for inclusion
in the database was largely based on the search terms
minimum & viable and extinction. Although other
terms were used, there was undoubtedly a bias towards
research in which MVP was an explicit consideration of
the original study, which implies an existing concern that
genetic issues may limit population persistence. However,
the database has missed many island taxa that have
survived for millennia with populations well below the
magic number of 5000 adult individuals [2]. Had these
very small island populations been included, the median
MVP might have been much lower. As with the Stewart
Island robin Petroica australis rakiura [4], the apparent
lack of inbreeding avoidance strategies [5] suggests there
must be either a reduced genetic load or fixed deleterious
alleles after population bottlenecks in these very small,
island populations. Such populations have a high level of
genetic fragility that limits their capacity to adapt, but it
does not mean their loss is inevitable. As noted recently
[6], assessments of population size need a palaeoecological
perspective. Such a perspective suggests modern parallels
with past events. For instance, most island taxa probably
had larger populations before sea levels rose at the start of
the Holocene, which caused populations to decline catastrophically before they successfully stabilised. This has
important implications beyond populations that are small
now. During the Pleistocene, many taxa survived in tiny
refuges and then flourished as the climate ameliorated [7].
Such taxa may thus be preselected to persist should their
populations again decline, and thus warrant ongoing conservation investment despite recent anthropogenic losses
that might have reduced their populations below 5000
individuals.
Even if the concept of an MVP were to be accepted as
useful, there are major issues with the data sets [3,8] used
to corroborate the theoretical calculations of MVP in [2].
First, although it would not be expected that all individual
examples fit a theoretical model, absurd results can be
removed. For instance, one study estimated the MVP of the
Lord Howe Island woodhen Tricholimnas sylvestris at 100
[9] (column H in the Appendix to [3]; and all credit to the
Corresponding author: Garnett, S.T. (stephen.garnett@cdu.edu.au).

618

authors for providing the data. In [3] this was standardised

as 126 (back-transformed from column I) to allow comparison between studies based on 40 generations and a 99%
probability of persistence. Given that the carrying capacity
of the island is estimated at 220, a figure unlikely to have
been exceeded in the last seven millennia, both estimates
seem reasonable. However, a second estimate of 2151 is
also presented for the same species. This is described as
a corrected MVP and is derived from an unpublished
Appendix to [8]. How a corrected MVP can exceed the
maximum evolutionary population size by an order of
magnitude is not explained. Second, some of the MVPs
in [3] are very odd when they are standardised. The most
extreme is an estimate of the MVP for the grizzly bear
Ursus arctos horribilis that was originally 101 individuals
(six other estimates ranged from 40 to 250); when standardised, this increases to 44 259 bears (which would need
an area equivalent to nearly 20% of the contiguous 48 USA
states). Third, and most importantly, it is unclear how the
much-quoted medians in [3] were derived. The median of
the original MVP estimates is only 375 if duplicate MVPs
for the same taxa are sampled randomly. However, the
median of the original MVPs seems to be approximately
the 3299 quoted in [3] because only corrected MVPs from
the unpublished Appendix to [8] are used when there are
duplicate MVPs.1 For standardised MVPs in [3], the median would be 1192 not 4169 were data other than those from
[8] given any credibility. This is because, on average, the
corrected MVPs in [8] are 26 times the standardised MVPs
derived from other studies of the same taxa (median 10.9)
and 94 times the original MVPs (median 30.8); the only
cases for which the corrected MVPs are lower than the
standardised MVPs are when the standardisation process
has, as with grizzly bears, greatly exaggerated the original
estimate. If the highly precautionary corrected MVPs
from [8] are removed entirely from the analysis, the median standardised MVP is only 742. The median nonstandardised MVP is just 250.
As a final point, it is certainly a good thing that Homo
sapiens is not in the early phase of its evolutionary history
and requiring conservation investment. One line of research suggests that the total human population
remained at approximately 1000 individuals for nearly
a million years [10]. Arguably, however, we are more
vulnerable to extinction due to anthropogenic causes
now than we were to any threat when our population
was 20% of the theoretical MVP. As concluded in [1],
1
Taxonomic ambiguity in [3] makes it difficult to see exactly which estimates are
considered duplicates, but this ambiguity is not enough to explain a 10-fold difference.

Update
although it is essential to manage genetic issues in conservation programs, the data from the studies assembled
in [3] do not justify the recommendation that taxa with
fewer than 5000 individuals should receive less conservation funding.
References
1 Flather, C.H. et al. (2011) Minimum viable populations: is there a
magic number for conservation practitioners? Trends. Ecol. Evol. 26,
307316
2 Traill, L.W. et al. (2010) Pragmatic population viability targets in a
rapidly changing world. Biol. Conserv. 143, 2834
3 Traill, L.W. et al. (2007) Minimum viable population size: a metaanalysis of 30 years of published estimates. Biol. Conserv. 139, 159166
4 Laws, R.J. and Jamieson, I.G. (2011) Is lack of evidence of inbreeding
depression in a threatened New Zealand robin indicative of reduced
genetic load? Anim. Conserv. 14, 4755

Trends in Ecology and Evolution December 2011, Vol. 26, No. 12

5 Jamieson, I.G. et al. (2009) Why some species of birds do not avoid
inbreeding: insights from New Zealand robins and saddlebacks. Behav.
Ecol. 20, 575584
6 Rull, V. and Vegas-Vilarrubia, T. (2010) What is long-term in ecology?
Trends. Ecol. Evol. 26, 34
7 de Bruyn, M. et al. (2011) Faunal histories from Holocene ancient DNA.
Trends Ecol. Evol. 26, 405413
8 Reed, D.H. et al. (2003) Estimates of minimum viable population sizes
for vertebrates and factors influencing those estimates. Biol. Conserv.
113, 2334
9 Brook, B.W. et al. (1997) How secure is the Lord Howe Island woodhen?.
A population viability analysis using VORTEX. Pac. Conserv. Biol. 3,
125133
10 Hawks, J. et al. (2000) Population bottlenecks and Pleistocene human
evolution. Mol. Biol. Evol. 17, 222
0169-5347/$ see front matter 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tree.2011.08.005 Trends in Ecology and Evolution, December 2011,
Vol. 26, No. 12

Letters

Minimum viable population size: not magic,

but necessary
Barry W. Brook1, Corey J.A. Bradshaw1,2, Lochran W. Traill3 and Richard Frankham4
1

Environment Institute and School of Earth and Environmental Sciences, University of Adelaide, Adelaide, SA 5005, Australia
South Australian Research and Development Institute, PO Box 120, Henley Beach, SA 5022, Australia
3
Imperial College London, Silwood Park, Berkshire, UK, SL5 7PY
4
Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia
2

We agree with Flather and colleagues [1] that there is no

magic number or universal threshold around which one
can plan for threatened species management to cover all
contingencies; neither have we ever claimed so [2]. As
Flather and colleagues reiterate [1], a minimum viable
population size [MVP; the abundance above which the
probability of extinction (over conservation-relevant timescales) is unacceptably low for any species] is illusory.
There is substantial variation in MVP among species [3]
and probably across subpopulations for widespread or
spatially disjunct species, and there is no obvious decision
threshold, as reviewed by Traill et al. [4] and elsewhere [5].
Yet even with this uncertainty, ignoring MVP because of
concerns over its imperfections or risk of misuse, as Flather
et al. [1] seem to prefer, would be imprudent.
First, we did not revive the MVP concept. It is a core
component, for instance, of the IUCN Red List of Threatened Species rules (e.g. Criterion D is based on minimum
stable abundances and Criterion C on a combination of
decline and population size; http://www.iucnredlist.org).
Second, there is a strong relationship between extinction
risk and population size that is supported by theory
(e.g. inbreeding and loss of genetic diversity, demographic
and environmental stochasticity, and extinction vortex),
simulation studies and substantial empirical data
(e.g. geneticdemographic experiments, island biogeography, and mammals on mountain tops) [38], especially
when appropriate generational scaling is used [9]. Third,
Corresponding author: Brook, B.W. (barry.brook@adelaide.edu.au).

Flather et al. [1] completely side-step the issue of genetic

erosion in small populations, and the substantial evidence
that inbreeding does indeed matter profoundly for extinction risk [7,10] (the genetic arguments alone are sufficient
to embrace MVP generalisations). They also object to our
methods for standardising MVP to account for intermodel
differences, varying life-history strategies and other scaling issues [4,5], but do not defend their implication that
not to attempt any standardisation is preferable.
Beyond the scientific and technical debates about MVP,
the broader issue raised by the Flather et al. review [1] is:
what to do if MVPs are ignored? The authors claim to: offer
suggestions for how conservationists might proceed in the
absence of such estimates but ultimately, provide little
more than: there is no substitute for diagnosing and
treating the mechanisms behind the decline of a population. We agree, in principle (who could not?). However,
there are serious limits to how much of this can be done in
most countries, or for most species [2]. Therefore, we must
rationally consider the alternatives as the conservation
crisis mounts. One is to rely on expert opinion, but evidence
shows that qualitative judgements and human intuition
are not reliable in this and other complex areas [11].
Another is to be guided by general principles that are
underpinned by theory, data and models, and which integrate multiple factors (including feedbacks and synergies),
treating uncertainty and assumptions explicitly and transparently. Model-based estimates of population viability are
analogous to other areas of complex system science, such as
weather forecasting, in that they are at least able to be
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