bleaching may ultimately help reef corals to
survive the recurrent and increasingly
severe warming events projected by current
climate models of the next half-century
3
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sacrifices short-term benefits for long-term
advantage. This counters conventional
wisdom that bleaching is detrimental from
all perspectives, and supports the role of
symbionts as adaptive agents
10,11
.
Andrew C. Baker
Wildlife Conservation Society, Osborn Laboratories
of Marine Science, New York Aquarium,
Surf Avenue at West 8th Street, Brooklyn,
New York 11224, USA
e-mail: abaker@wcs.org
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2850–2853 (1995).
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Thesis, Univ. Miami (1999).
9. Toller, W. W., Rowan, R. G. & Knowlton, N. Biol. Bull. Mar.
Biol. Lab. W oods Hole (in the press).
10. Buddemeier, R. W. & Fautin, D. G. Bioscience 43,
320–326 (1993).
11. Rowan, R. J. Phycol. 34, 407–417 (1998).
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Biol. Bull. Mar. Biol. Lab. W oods Hole 200, 51–58 (2001).
13. Buddemeier, R. W. & Smith, S. V. Am. Zool. 39, 1–9 (1999).
14.Wilkinson, C. et al. Ambio 28, 188–196 (1999).
Population control
African elephants and
contraception
P
rotected from hunting and provided
with access to water-holes during
droughts, elephant numbers can dou-
ble in a decade, severely damaging natural
vegetation and the many species dependent
upon it. Culling is an effective but contro-
versial control strategy, so Fayrer-Hosken et
al.
1
have assessed the efficacy of using
immunocontraception through vaccina-
tion, concluding that this could be a practi-
cal way of controlling elephant numbers.
However, an intervention feasible in repro-
ductive physiology may not be a practical
way to control a population. Fayrer-Hosken
et al. have not considered calculations
2,3
that
undermine the practicality of their method,
nor alternative management strategies.
Controlling elephants in Kruger
National Park, South Africa, by immuno-
contraception would necessitate treatment
of 2,250 cows each year over an initial
period of 11 years (ref. 3). Even if individual
treatments were 100% effective, the costs
would be likely to exceed the total manage-
ment budget of the South African national
parks. The best results of Fayrer-Hosken et
al. involved two of ten elephants becoming
pregnant, and that was after receiving two
booster vaccinations.
The effectiveness of this method may be
less than claimed. Of the control group,
89% became pregnant within a year. This
seems high, exaggerating the difference
between treated and control groups. Data
from 813 adult cows culled in Kruger
National Park between 1979 and 1994
showed that 51% (range, 36–77%) were
pregnant. This is to be expected: gestation
lasts 22 months and the calving interval is
44 months (ref. 2), so about 50% of a
sample of cows should be pregnant. Thus,
on average, females go for 22 months with-
out becoming pregnant. In a random sam-
ple of females monitored for 12 months,
only 55% (not 89%) should therefore
become pregnant.
Between 16 and 1,846 elephants of all
age classes and both sexes were culled annu-
ally in Kruger National Park from 1967 to
1994. We share the desire to reduce culling
and have sought methods to do so. Remov-
ing or sterilizing 250 subadult females each
year should reduce population growth to
zero
2,3
. Moreover, densities of greater than
0.37 elephants per square kilometre result
in reduced population growth rates —
probably due to reduced reproductive out-
put by newly sexually matured females or to
increased calving intervals
2
. Culling, as
conducted, maintained densities at which
population growth was near its maximum.
Culls should be delayed for one year after
counts exceed 0.37 elephants per square
kilometre to allow density dependence to
reduce numbers naturally
2
. Culls may still
be necessary, but they would then be much
less frequent and involve far fewer animals.
Stuart L. Pimm*, Rudi J. van Aarde†
*Center for Environmental Research and
Conservation, Columbia University,
1200 Amsterdam Avenue, New York,
New York 10027, USA
e-mail: stuartpimm@aol.com
†Conservation Ecology Research Unit,
University of Pretoria, Pretoria 0002, South Africa
1. Fayrer-Hosken, R. A., Grobler, D., Van Altena, J. J.,
Bertschinger, H. J. & Kirkpatrick, J. F. Nature 407, 149 (2000).
2. van Aarde, R. J., Whyte, I. & Pimm, S. Anim. Conserv. 2,
287–294 (1999).
3. Whyte, I., van Aarde, R. J. & Pimm, S. Anim. Conserv. 1,
77–83 (1998).
Fayrer-Hosken et al. reply –– Pimm and van
Aarde question the feasibility of controlling
elephant numbers by immunocontracep-
tion, arguing that the sterilization or
removal of 250 subadult cows each year is
the answer to population growth. However,
there are no known safe methods of
sterilizing free-roaming African elephants.
Moving 250 subadult females to another
park is impractical as there are very few
areas able to receive elephants from Kruger
National Park without becoming confront-
ed with an elephant overpopulation prob-
lem of their own.
This number of subadult cows cannot
be moved without disrupting the social
order within their herds. Keeping them in
their herds would mean that (assuming a
mean herd size of 12.4, as shown in our
study, and an average of 3 subadult females
per herd) Kruger National Park would have
to move 1,033 elephants — an unrealistic
and expensive proposition. Hence the only
practical way to remove 250 subadult
females would be to cull them, which
Pimm and van Aarde agree is an unaccept-
able solution.
We have shown that immunocontracep-
tion using porcine zona pellucida (pZP)
works in the African elephant, although its
long-term effectiveness in controlling pop-
ulations is still being evaluated in South
Africa. The cost and speed of field delivery
have not been assessed for vaccinating large
groups of elephants. However, contrary to
the calculations of population modellers
1,2
,
immunocontraception has worked in herds
of wild horses and white-tailed deer
3
.
Preserving these magnificent creatures
and their genetic contribution for the
future is a common goal. On the basis of a
single administration of a multiple-release
pZP vaccine that is being developed for use
in horses (I. K. M. Liu, personal communi-
cation), it should be possible to reduce the
first three vaccinations used in our original
study to a single dose and so minimize the
stress, cost and labour of elephant
immunocontraception.
We therefore question Pimm and van
Aarde’s criticism regarding the practicality
of field immunocontraception for Kruger
Park’s elephant herds. It is our judgement
that preserving these animals through
immunocontraception is a realistic strategy
that would save elephants without having
to kill them.
R. A. Fayrer-Hosken*, D. Grobler, J. J. van
Altena, H. J. Bertschinger, J. F. Kirkpatrick
*College of Veterinary Medicine, University of
Georgia, Athens, Georgia 30602-7385, USA
e-mail: rfh@calc.vet.uga.edu
1. van Aarde, R. J., Whyte, I. & Pimm, S. Anim. Conserv. 2,
287–294 (1999).
2. Whyte, I., van Aarde, R. J. & Pimm, S. Anim. Conserv. 1,
77–83 (1998).
3. Kirkpatrick, J. F., Turner, J. W., Liu, I. K. M., Fayrer-Hosken,
R. A. & Rutberg, A. T. Reprod. Fertil. Dev. 9, 105–110 (1997).
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