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Wildlife Research
Volume 26, 1999
CSIRO Australia 1999

A journal for the publication of original scientific research

in the biology and management of wild native or feral introduced vertebrates

w w w. p u b l i s h . c s i r o . a u / j o u r n a l s / w r
All enquiries and manuscripts should be directed to
Wildlife Research
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for CSIRO Australia and
the Australian Academy of Science

q CSIRO 1999

Wildl. Res., 1999, 26, 227237


Analysis of the impact of stoats, Mustela erminea,

on northern brown kiwi, Apteryx mantelli,
in New Zealand
B. BasseA , J. A. McLennanB and G. C. WakeA

Mathematics and Statistics Department, University of Canterbury,

Private Bag 4800, Christchurch, New Zealand.
Landcare Research, Private Bag 1401, Havelock North, New Zealand.

An age-structured population analysis is used to determine recruitment levels and a condition for
survival which can assist management decisions and hence improve the viability of populations of
northern brown kiwi, Apteryx mantelli, in forests on the New Zealand mainland. Currently, in the
absence of predator control, recruitment rates are less than 5% due to high levels of stoat, Mustela
erminea, predation on juvenile kiwi. Predation levels on adult kiwi are very low. The analysis predicts
that a recruitment rate of 19% is required to maintain population stability. To achieve this target, stoat
populations have to be reduced by about 80% in some years, and maintained at a critical residual
density estimated to be a value less than two animals per square kilometre for up to nine months until
immature kiwi reach a safe size of about 1200 g (50% of their adult weight). Recent predator-control
initiatives indicate that stoat numbers can be reduced and maintained at low levels in relatively small
areas of mainland forest (up to 1000 ha). New techniques are needed to protect kiwi over larger areas.

All four species of kiwi (Apterygiformes) have declined significantly in range and
abundance since human settlement. Little spotted kiwi (Apteryx owenii Gould) are now
probably extinct on the mainland; great spotted kiwi (Apteryx haastii Potts) are now mainly
confined to high-rainfall regions of the north-western corner of the South Island; northern
brown kiwi (Apteryx mantelli Bartlett) have disappeared from the lower third of the North
Island; and southern brown kiwi (Apteryx australis Shaw & Nodder) are now largely
restricted to Fiordland and Stewart Island.
Over the last century, kiwi abundance has probably declined by at least 90% in most
North Island forests. Bullers (1887, 1888) historical accounts suggest former densities of
40100 adults km2 , whereas present densities seldom exceed 4 adults km2 (McLennan and
Potter 1992). Some parts of Northland and two offshore islands (Kapiti and Little Barrier)
still support 50100 adults km2 . Mainland populations, however, contain significantly
fewer juveniles (McLennan and Potter 1993) than do similar populations on predator-free
offshore islands (Colbourne 1992).
Kiwi evolved in the absence of mammalian predators, but co-exist today with as many
as seven obligate and facultative carnivores introduced by Polynesians and Europeans
(Wodzicki 1950). Recent studies show that a range of predators eat kiwi and their eggs,
with different species being involved at different stages (McLennan et al. 1996). Impacts
of predators on adults and eggs vary from place to place, but are often small and of
little consequence. Fortunately, adult brown kiwi (23 kg) exceed the threshold prey size
of cats and stoats, the largest of the common predators in mainland forests. Similarly,
kiwi eggs benefit from their large size in that they are too big (mean length 129 mm)



B. Basse et al.

and heavy (435 g) to be punctured or removed by rats. Immature kiwi, on the other
hand, consistently suffer heavy losses to stoats (Mustela erminea) and too few survive to
replace adult deaths in most years. This is the main cause of kiwi decline in mainland
forests (McLennan et al. 1996), though other factors (e.g. accidental poisoning in possum
(Trichosurus vulpecula Kerr) control operations) accelerate the process in some areas.
Stoats were first introduced to New Zealand in 1855 to help control rabbits (Oryctolagus
cuniculus) (Wodzicki 1950). They dispersed rapidly, aided by repeated liberations, and had
probably spread throughout both the North and South Islands by the turn of the century
(King 1990). Kiwi and stoats have therefore co-existed in mainland forests for approximately
100150 years. Stoat abundance varies between localities and years, apparently in response
to changes in the availability of their main foods (birds, rodents, lizards and invertebrates).
These changes are particularly pronounced in beech (Nothofagus spp.) forests, where heavy
seeding is followed by an irruption of mice, then stoats (King 1983). Measurements suggest
that even when stoats are at the low point of their cycle, they are still sufficiently abundant
to prevent adequate recruitment of kiwi (McLennan et al. 1996). Other less-vulnerable
species such as yellowhead (Mohoua ochrocephala) get a reprieve during low-stoat years
and make good some of the losses incurred during an irruption (ODonnell et al. 1992).
Here we identify: (1) the recruitment and predation rates needed to maintain kiwi
populations in mainland forests, and (2) the critical density that stoats cannot exceed to
ensure the desired recruitment rate is achieved.
Age-structure analysis of the population
The population is divided into two age classes, immature kiwi (differentiated by chicks and
juveniles) and adults (differentiated by gender) (Fig. 1). Chicks develop into independent juveniles
when they leave the nest permanently, approximately 1825 days after hatching (McLennan 1988).
Juveniles become adults at an age of about 18 months (Reid and Williams 1975).

Fig. 1.

Schematic diagram showing the age compartments of the population.

In particular, we are interested in the death rates due to predators of immature kiwi. If 1 and 2
are the (constant) natural mortality rate of chicks and juvenile kiwi per capita per year respectively
and 1 (s/S) and 2 (s/S) are the (constant) mortality rate of chicks and juvenile kiwi per capita per
year due to predators then the rates of chicks and juvenile kiwi leaving the cohort due to death are
(1 + 1 (s/S)) and (2 + 2 (s/S)) per capita per year respectively.

Impact of stoats on northern brown kiwi


We need to further clarify the predation rate (s/S) and the fraction part (s/S). Since stoats are
the main predators of immature kiwis (McLennan et al. 1996), we assume that the effect of other
predators, besides stoats, on immature kiwi is negligible. The predation rate of immature kiwi per
capita per year is (s/S) where s is the stoat density per square kilometre at time t (measured in
years) and S is the maximum density of stoats per square kilometre. (In practical terms, S is the
maximum carrying capacity of stoats in any New Zealand forest.) The fraction (s/S) is a number
between 0 and 1 and (s/S) represents a predation rate that ranges between 0 (when there are no
stoats, i.e. s = 0) and (when s = S, that is, when stoats are at maximum density.) The scaling
factor, (s/S), is important (as opposed to having a single parameter to represent mortality) because
we want to identify the critical density sc at which (sc /S) (the linear combination of the predation
rates for chicks and juveniles) is low enough to allow kiwi to survive.
Let n 1 (t) and n 2 (t) be the number of male and female adult kiwi respectively at time t in years.
Let f 1 and f 2 be the respective male and female mortality rates. Since adult kiwi are too large to be
killed by stoats, we treated their mortality rates as a constant. If b is the productivity rate per adult
female per year then the number of chicks hatched in one year is given by bn 2 (t).
For survival, a female adult kiwi must produce at least one female chick that survives to adulthood.
That is, each female must replace herself. We will use this principle to derive a survival threshold (a
critical predation rate) at which juvenile recruitment equals adult mortality.
The parameters and variables used in the analysis are summarised in Table 1.
Table 1.

Summary of parameters and variables

The age (in years) when a chick becomes a juvenile
The age (in years) when a juvenile becomes an adult
The number of adult males at time t
The number of adult females at time t
Number of young produced per adult female per year
Natural death rate of chicks per capita per year
Natural death rate of juvenile kiwi per capita per year
Predation rate of chicks per capita per year
Predation rate of juvenile kiwi per capita per year
Death rate of adult males per capita per year
Death rate of adult females per capita per year
The stoat density per km2 present at time t
The maximum possible density of stoats per km2
The critical stoat density per km2 required for survival of kiwi


n 1 (t)
n 2 (t)
1 (s/S)
2 (s/S)

0 06
0 85
4 76
0 365
3 97
0 082
0 082

The data set and parameter values used in the analysis

The values for the parameters used in this analysis were taken from the data set compiled by
McLennan et al. (1996), the largest available for any species of kiwi. The productivity estimate of
b = 0 85 chicks per female per year was derived from 33 pairs in three different areas. The sex
ratio of adults in North Island forests varies from place to place but has no consistent bias, implying
that average mortality rates are similar for males and females; thus, the adult mortality estimate of
f 1 = f 2 = 0 082 per capita per year was based on 122 1 radio-tracking-years of 171 individuals in
nine locations.
Mortality estimates of chicks (1 = 4 76, and 1 (s/S) = 3 97 per capita per year) and juvenile
kiwi (2 = 0 365 and 2 (s/S) = 3 1 per capita per year) were derived by determining the fates of
42 individuals in four locations. Deaths of immature kiwi were attributed to predators (64%) or other
causes (33%), according to criteria listed in McLennan et al. (1996).
Estimates of stoat densities in New Zealand forests were derived from: relative abundance indices
(King 1983; Murphy and Dowding 1995); counts of animals removed during intensive kill-trapping
operations (Murphy and Bradfield 1992; McLennan 1997); mark and recapture studies (Murphy and
Dowding 1995); measurements of territory size and territory overlap (Murphy and Dowding 1994;
1995); and biomass values for various predators in New Zealand forests (Brockie 1992).


B. Basse et al.

In beech forests following heavy seeding, stoats appear to reach densities of 79 resident animals
km2 . These estimates are based on a mark and recapture study (12 animals per 150 ha) and
measures of the size of exclusive, core territories (20 ha for females, 25 ha for males). They do not
include non-territorial and dispersing animals, so are conservative. Nevertheless, they are very similar
to McLennans (1997) estimate of 9 animals km2 , derived by intensive kill-trapping on a 750-ha
peninsula with restricted opportunities for immigration (65 animals removed over 3 months). We
therefore assume that stoat densities in New Zealand forests reach a maximum of S = 10 animals
km2 .
Trapping indices suggest that stoat densities in beech forests vary five-fold between years, depending
on the stage of the beech seeding-cycle (King 1983; Murphy and Dowding 1995). Natural fluctuations
in stoat density therefore appear to be within the range of 210 animals km2 . Densities may be
lower in forests where cats are plentiful (Brockie 1992) although, in such areas, the combined biomass
of carnivores (0 032 kg ha1 ) is equivalent to a stoat density of 1011 adults km2 .

Finding the critical recruitment rate required for survival
Assuming no stochasticity, the number of chicks that reach adulthood, is:

bn 2 (t 15)e(+(s/S))1 5

juvenile natural mortality
chick natural mortality
per year
per year

(0061 + 1442 ) = 05 per capita per year

juvenile predation rate
chick predation rate
per year
per year

0061 + 1442
= 31 per capita per year

and n 2 (t 15) is the number of females in the kiwi population at time (t 15), which
is when the juveniles reaching adulthood were hatched. bn 2 (t 15) is the number of
chicks hatched at time (t 15) and e(+(s/S))1 5 represents the exponential decline
(since immature kiwis leave the cohort at a rate proportional to numbers present) due to
the combined natural and predator rates of the chicks and juveniles.
Finding the threshold for survival
It is possible to find a series expansion for the number of adult males and females
explicitly and hence find out the asymptotic behaviour of the adults (Basse et al. 1997)
as er t where r is explicitly determined and r = 0 corresponds to the threshold between
survival and extinction.
Here we show that the criterion for survival of female kiwi can be derived using the
principle that each female must, in her lifetime, produce on average at least one female
chick that reaches maturity. A female has an average life expectancy of (1/ f 2 ) years. Of
the chicks hatched in her lifetime, (1/ f 2 ) (b/2) e(+(s/S))1 5 survive 1 5 years to
become female adults. Thus, a survival condition for the whole kiwi population is:

b (+(s/S))1 5
2 f2

Impact of stoats on northern brown kiwi


The above inequality can be rearranged to give


2 f2

2 0082
= 06 per capita per year.
This gives a critical value for the annual predation rate of immature kiwi of (sc /S) = 06
per capita per year. If the predation rate (s/S) falls below the critical value (sc /S) = 06
per capita per year then this will enable the population to sustain itself, while a predation
rate above the critical value will eventually lead to extinction. Our criterion for survival
is therefore:


The actual value of (s/S) derived from field measurements is (s/S) = 31. This
greatly exceeds the critical value (sc /S) = 06, and has to be reduced by a factor of
five to ensure kiwi survival (Fig. 2).

Fig. 2.

Immature recruitment versus predator mortality.


B. Basse et al.

Fig. 3. Sensitivity analysis of (sc /S) = (1/1 5) ln((b/2 f 2 )) with = 0 5. (a) (sc /S)
as a one-dimensional function of f 2 (with b = 0 85) and b (with f 2 = 0 082) respectively.
(b) (sc /S) = (1/1 5) ln((b/2 f 2 )) as a two-dimensional function (with only positive function
values plotted) of f 2 and b. The actual parameter values used in the modelling were b = 0 85 and
f 2 = 0 082.

Recall that the number of chicks hatched that reach adulthood is

bn 2 (t 15)e(+(s/S))1 5 .
In our data set, the number of kiwi hatched at time (t 15) was bn 2 (t 15) = 42.
Given the threshold value of (sc /S) = 06, the number that need to be recruited to
maintain population stability is 42e(0 5+0 6)1 5 8 or 19% of those hatched. The
population can therefore sustain an 81% loss of juveniles without declining. In reality,
the survival of immature kiwi is 42e(0 5+3 1)1 5 0. Clearly, the losses to stoats are
threatening the population with extinction.

Impact of stoats on northern brown kiwi


Sensitivity analysis of the threshold condition

The critical value for the predation rate per capita per year of immature kiwi is

2 f2

It is interesting to see how this critical value changes with small changes in either b or
f 2 . It should be noted that is just a linear term so (sc /S) changes linearly with
changes in . Since all parameters must be non-negative the feasible region for b and f 2
is b 2 f 2 e1 5 . If b is close to zero then the critical value is extremely sensitive to
small changes in b but b = 0 is not in the feasible region. Fig. 3 shows graphically
(sc /S) as a one- and two-dimensional function of b and f 2 . Note that from McLennan
et al. (1996), b can be in the range 01 72 per year and f 2 is 0 040 14 per year.
Reducing the stoat population to find the threshold for survival
We have assumed that stoats are the main predator of young kiwi, and that reducing
the stoat population is the only way of reducing the predation rate. We also assume that
predation losses change in direct proportion to stoat abundance. Thus, stoat populations
have to be reduced by 80% in order to reduce predation losses by 80%. Curve a in Fig. 4
suggests that the survival threshold (19% recruitment) is reached when stoats decline to
a density of 1 94 animals km2 . However, this critical density estimate is conservative
because it assumes that stoats were at maximum density when the survival rates of immature
kiwi were measured in the field. In fact, the data were collected over a range of sites
and years, and so reflect average losses over an unknown range of stoat densities. If,

Fig. 4.

Immature recruitment versus stoat density.


B. Basse et al.

at the time of the measurements, stoats were at an average density of 6 rather than 10
animals km2 (see curve b in Fig. 4) then the critical density for kiwi survival would be
calculated as 1 16 animals km2 rather than 1 94 km2 . We cannot find the exact value
for the critical stoat density but we know it is between 0 and 2 animals km2 . Thus, to
ensure kiwi survival, stoat populations have to be reduced by at least 80% in years when
they are abundant.
Population declines of kiwi
Our results support the view of McLennan et al. (1996) that northern brown kiwi are
failing in mainland forests because of predation by stoats on immature kiwi. Current survival
rates of juveniles (15%) are well below the threshold required to maintain populations
(19%). At the present rate of loss (about 6% per annum) many of the remaining populations
in the North Island will disappear over the next 4050 years. Some have already done so
(McLennan and Potter 1992; Miller and Pierce 1995). The estimate of time to extinction
is an approximation, based on the assumptions that the rate of loss is constant in time
and space, that demographic stochasticity and other chance events cause extinction when
densities fall below 0 2 individuals km2 , and that there are currently 2 5 kiwi km2 in
occupied forests.
The first assumption is clearly not true, though it makes no difference to our estimate.
Residual kiwi densities vary considerably within and between regions, and simple back
projections show that average rates of loss over the last 100 years must have been less
than 6% in areas that currently support 1 or more kiwis km2 . In fact, it appears likely
that the decline started slowly, then accelerated as the birds became rarer. This pattern
would be expected if predation losses diminish at high kiwi densities (Pech et al. 1995), a
possibility now being investigated. King (1984) argued that predators are now unlikely to
cause further avian extinctions in mainland habitats of New Zealand because the vulnerable
species have already gone. However, she listed five species, all ancient endemics with
restricted distributions, that might be declining because of predation, and commented that
other at-risk species could be identified following further research. King (1984) considered
that sufficient time had elapsed for the full impacts of introduced predators to become
apparent, a view that is not supported by our results or those of other recent investigations.
All of the five species on Kings (1984) list have continued to decline over the intervening
decade, with predation being confirmed as the main cause in at least three cases: kokako
(Callaeas cinerea), black stilt (Himantopus novaezealandiae), kakapo (Strigops habroptilus).
Other species now known or suspected to be declining because of predation include
yellowhead (Mohoua ochrocephala) (ODonnell et al. 1992), kaka (Nestor meridionalis)
(Beggs and Wilson 1991), northern brown kiwi (Apteryx mantelli) and great spotted kiwi
(Apteryx haastii) (McLennan et al. 1996), New Zealand dotterel (Charadrius obscurus)
(Dowding and Murphy 1993) and North Island weka (Gallirallus australis) (Bramley 1996).
Remarkably, New Zealand pigeon (Hemiphaga novaeseelandiae), a relatively common species
by todays standards, also appears to be declining because of predation and (possibly)
competition from introduced mammals (Clout et al. 1995).
Other species currently regarded as being safe may well be in a similar predicament,
but few have been studied in sufficient detail to determine the factors regulating their
distribution and abundance.
Clearly, predator impacts are far from over in mainland habitats of New Zealand. Species
that are still relatively common have not demonstrated an ability to co-exist indefinitely
with introduced mammals. Predator-induced declines can take much longer than 150 years
to run their course. Certainly, the most vulnerable species have already gone, presumably
because predator impacts were severe at all stages of their life cycle. Other species, such
as northern brown kiwi, have persisted for longer because predation impacts are either

Impact of stoats on northern brown kiwi


intermittent, restricted to one stage of the life cycle, restricted to particular habitats, or
restricted to one sex. Nevertheless, they are declining, but are just doing it more slowly.
Critical stoat density
Our analysis predicts that the critical stoat density, at which recruitment of juvenile kiwi
balances adult losses, is estimated to be a value less than 2 animals km2 . To achieve
this density, stoat populations have to be reduced by at least 80% in years when they are
abundant. These management targets are tentative estimates based on limited measurements
of stoat density in New Zealand forests and the assumption that predation losses of young
kiwi are directly proportional to stoat abundance and independent of kiwi density (that is,
it is a type I functional response). Nevertheless, two predictions of the analysis appear
to be realistic: (1) that stoat populations have to be reduced substantially in order to
produce any measurable benefits, and (2) that in most habitats, stoat densities seldom, if
ever, decline naturally to levels that allow adequate recruitment in kiwi.
The shape of each recruitment curve in Fig. 4 is such that small changes in stoat
density of 05 animals km2 greatly affect survival rates of juvenile kiwi. Once stoat
densities reach 5 animals km2 , further increases are largely irrelevant, because by then
chick survival rates are already close to zero. In plague years, for example, the removal
of 50% of the animals would have no significant effect on juvenile survival. The analysis
therefore predicts that predation rates will not necessarily decline following the removal
of large numbers of predators the outcome of many mustelid-control operations in New
Zealand (e.g. Reid et al. 1995). Indeed, if trapping operations generally remove only about
50% of the stoats present, as (King and McMillan 1982) suggest, most will fail to reduce
predation rates on young kiwi.
Trapping indices (captures per 100 trap-nights) suggest that densities of stoats in beech
(Nothofagus) forests vary about five-fold between years (King 1983; Murphy and Dowding
1995). Densities are both higher on average and less variable in mixed podocarp forests
and agricultural landscapes where food supplies are more constant (King 1990). These
natural variations in density appear to be within the range of 210 animals km2 , and thus
seldom benefit immature kiwi. It is noteworthy, however, that kiwi are declining more
slowly in sub-alpine regions of the South Island than elsewhere (McLennan and McCann
1994), possibly because in harsh, upland habitats stoats sometimes dip below the critical
density required for adequate recruitment.
Stoat densities vary seasonally within habitats, with a peak in midsummer. Mortality
of both pre-independent and newly-independent young is always very high, so the peak is
short-lived (King and McMillan 1982). Nevertheless, stoats are most plentiful in forests
when young kiwi have either just become independent or are about to do so. At these
times, they outnumber kiwi chicks by as much as ten to one. Each stoat therefore has a
limited opportunity to kill an immature kiwi, and most of them probably never do. Young
kiwi are merely an incidental item in their diet, a rare but nevertheless highly-vulnerable
and easily-captured prey species. The general absence of kiwi remains in the digestive
tract of stoats is therefore an expected result (King and Moody 1982; King 1983; Murphy
and Dowding 1995), but one that belies the true relationship between the two species.
Significance of other predators
Our assumption that stoats are responsible for all predation losses of immature kiwi is
unlikely to hold in all habitats. Stoats are the most widely distributed carnivore in New
Zealand (King 1990), and are often the only large predator in forests dominated by beech
(Nothofagus). Predator communities are more diverse in edge habitats where rabbits are
present, and in forest tracts containing mixed hardwood and softwood assemblages. Some
parts of the North and South Islands support sympatric populations of feral cats (Felis


B. Basse et al.

catus), ferrets (Mustela furo), weasels (Mustela nivalis vulgaris) and stoats, in various
proportions depending on habitat type (King 1990). All four species kill immature kiwi.
Our analysis may be applicable to mixed predator assemblages, using combined densities
or biomass values, though we have not tested this. As it stands, it is applicable to all
large forest tracts in both the North and South Islands, where stoats are invariably the
dominant predator. It is probably of little or no use in small patches bordering farmland or
river-beds, where stoats are partially displaced by cats and ferrets. However, kiwi are now
rare in such habitats, except in parts of Northland and Taranaki (Reid and Williams 1975).
Future of kiwi in mainland forests
At present, immature kiwi are secondary prey and suffer excessive predation (Pech et al.
1995). They are picked off easily by introduced predators, whose numbers are maintained
at relatively high levels by other more common and less vulnerable prey species. As our
analysis indicates, the task of increasing recruitment rates of kiwi in mainland forests to
required levels is a daunting one, involving the removal of a high proportion of predators
from large areas over a long time. Current methods of stoat control (intensive trapping) are
only occasionally effective (e.g. McLennan 1997) and too expensive and labour-intensive
to apply on a large scale (>1000 ha). Clearly, the persistence of kiwi on mainland New
Zealand is now largely dependent on the development of new techniques for controlling
We thank Bill Lee and Tony Sinclair for commenting on the paper. We also thank the
numerous people who collected the data on kiwi upon which this study is based. This
study was funded by a grant from the New Zealand Foundation of Research, Science and
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Impact of stoats on northern brown kiwi


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Manuscript received 4 July 1997; accepted 2 July 1998