CSIRO PUBLISHING

www.publish.csiro.au/journals/wr Wildlife Research, 2008, 35, 417424

Demographic indications of decline in the spectacled flying fox

(Pteropus conspicillatus) on the Atherton Tablelands of
northern Queensland

Samantha Fox A,B,E,F, Jon Luly B, Catlin Mitchell B, Jenny MacleanC and David A. Westcott D
A
School of Marine and Tropical Biology, James Cook University, Townsville, Qld 4811, Australia.
B
School of Earth and Environmental Sciences, James Cook University, Townsville, Qld 4811, Australia.
C
Tolga Bat Rescue and Research, PO Box 685, Atherton, Qld 4883, Australia.
D
CSIRO Sustainable Ecosystems, Atherton, Qld 4883, Australia.
E
Present address: Wildlife Management Branch, Department of Primary Industries and Water,
GPO Box 44, Hobart, Tas. 7001, Australia.
F
Corresponding author. Email: samantha.fox@dpiw.tas.gov.au

Abstract. A lack of information about the spectacled flying fox (Pteropus conspicillatus) makes management and con-
servation of this vulnerable species difficult. The analysis of population dynamics using life-history traits and life tables
is widely used in planning for the conservation and management of wildlife. In the present study, the first life table for any
species of bat is provided and age estimates derived from counts of annual increments in tooth cementum rings are used
to assess population trends and life-history traits in the spectacled flying fox on the Atherton Tablelands in north
Queensland. As a result of high mortality, longevity was much shorter than expected from a theoretical basis. Life-table
analyses suggest that the population experienced a 16% decrease during the 2 years of study. Absence of extended
longevity to compensate for low reproductive output and delayed sexual maturity in slow end mammal species such as
P. conspicillatus reduces the window of opportunity for females to reproduce and adapt to changes in mortality rates. This
study suggests that spectacled flying fox populations are sensitive to increased mortality and that reducing mortality rates
should be the primary goal in conservation planning for P. conspicillatus.

Introduction
Life-history strategy is an important determinant of population
resilience because the potential rate of population growth is
intrinsically linked to reproductive capacity (Krebs 2001).
Thus, life-history studies can be informative in understanding
why some species suffer from a population decline while others
do not. Mammals with a slow life history are most at risk of
population declines and extinction (Purvis et al. 2000; Jones
et al. 2003) owing to the late age of their first reproductive
event and their low reproductive output. The projected popu-
lation growth in slow species is most sensitive to changes in
juvenile and adult survival rates (Oli 2004); therefore, mortal-
ity in those life stages is a critical variable in modelling poten-
tial extinction risk.
Larger mammals are often typified by having a slow life
history, whereas small species usually have a fast life history
(Stearns 1992). This fastslow continuum may persist,
however, even allowing for allometric differences (Partridge
and Harvey 1988; Purvis and Harvey 1995). For example,
despite their small size, bats demonstrate a slow life-history
strategy. The extraordinary longevity in bats is well known
(Austad and Fischer 1991; Wilkinson and South 2002; Barclay
and Harder 2003; Brunet-Rossinni and Austad 2004), with
members of the order Chiroptera living up to three times longer
than non-flying eutherian mammals of comparable size
(Austad and Fischer1991). The trade-off against this extended
life span is a low reproductive output (Barclay and Harder
2003; Barclay et al. 2004). Within the suborder Megachiro-
ptera, flying foxes in the genus Pteropus produce a single pup
annually (Hall and Richards 2000; Barclay and Harder 2003)
and make substantial maternal investment in its survival to
weaning (Hall and Richards 2000). Flying foxes have lived for
up to 30 years in captivity but there are few records of how long
they live in the wild (Hall and Richards 2000; Divljan et al.
2006) and like most bat species, population age structure is
unknown. Because of their high vagility, nocturnal habits and
tendency to roost in inaccessible places, demographic and life-
history data on bats remain scarce and no life tables have been
constructed for the group.
Although the low reproductive rate in flying foxes is
supposedly compensated for by extended longevity, it is also
one of two main predictors of extinction risk in this group (Jones
et al. 2003). More than half of the species within the genus
Pteropus are listed in the 2006 IUCN red list as threatened, with
seven species listed as critically endangered and four considered
extinct. Many species of flying fox, especially those found on
small Pacific Ocean islands, are facing imminent extinction
through habitat loss and subsistence hunting (Graham 1992;
Cousins and Compton 2005; Esselstyn et al. 2006).
In Australia, the spectacled flying fox (Pteropus conspicilla-
tus Gould, 1850) is a significant component of World Heritage

CSIRO 2008 10.1071/WR07127 1035-3712/08/050417

418 Wildlife Research
values in the Wet Tropics World Heritage Area (Westcott et al.
2001), and is listed as vulnerable under the Commonwealth
Governments Environment Protection and Biodiversity
Conservation Act 1999. Conservation and management of this
species is made all the more complex by the pest species status
accorded it by fruit farmers, and the previous practice of issuing
mitigation permits to cull marauding flying foxes. The specta-
cled flying fox contends with the same threats faced by all flying
fox species in Australia, such as habitat loss, electrocution on
powerlines, trapping on barbed wire fences, culling by fruit
farmers and colony disturbance, under permit or not. Since the
1980s, Atherton Tableland colonies of the spectacled flying fox
have also been subject to fatal envenomation by the paralysis
tick (Ixodes holocyclus). Some colonies can lose up to 5% of
animals in bad years (J. Maclean, unpubl. data).
The availability of large numbers of dead spectacled flying
foxes collected from a colony near Atherton enabled us to inves-
tigate the age structure of bats in the focal colony and to con-
struct life tables for the species. This is the first life table analysis
carried out on a wild flying fox population because of the logis-
tical and ethical difficulty of adequately sampling animals from
this group. Here, we develop life tables and estimate the mortal-
ity rate and intrinsic capacity for increase in P. conspicillatus. We
investigate basic life history parameters in a flying fox species,
ascertain whether traits and trade-offs conform to a theoretical
slow life history and examine how mortality rates influence the
dynamics of a wild flying fox population.
Papua
New Guinea
10
12
14
Cooktown
16 Wet Tropics World
Heritage Area
Tolga Scrub Cairns
colony
18
Townsville
20
22
140 142 144 146 148 150
S. Fox et al.
Methods
Data collection
Teeth were taken from 361 spectacled flying foxes (female = 196,
male = 165) that had died from tick paralysis at the Tolga Scrub
colony on the Atherton Tablelands in northern Queensland (Fig.
1) during 2001 and 2002. These animals were collected freshly
dead or had died after being taken into care. Tick paralysis
season for flying foxes corresponds with the time of year when
females give birth: OctoberDecember. Spectacled flying foxes
are seasonal breeders and, because we collected teeth at the
same time as parturition, all animals used in this study had just
had another birthday an important consideration when aging
an animal using an annually incremental feature. We recorded
sex, reproductive status and morphological measurements and
each individual flying fox was given a numerical identifier.
Female reproductive status was recorded in three categories:
(1) lactating at the time, had given birth that season; (2) had
given birth in previous years but not lactating at time of death;
or (3) had never suckled young. The difference between a
female that had lactated previously and one that had never
lactated was obvious by the elongation of the nipples caused by
offspring remaining attached for several months.
Ages of individual animals were estimated by counting
cementum layers around the root of the canine tooth, based on
the methods of Cool et al. (1994). Tooth cementum layers
provide an accurate representation of the age of captive
Fig. 1. Map of north Queensland indicating the
distribution of the spectacled flying fox, Pteropus
conspicillatus, in Australia, within the Wet Tropics
World Heritage area (grey shaded areas). The
location of the colony studied is indicated by the
open circle.
152
Indications of decline in the spectacled flying fox Wildlife Research 419

Australian flying foxes (Cool et al. 1994), a conclusion sup- estimates (mortality rate at the time when sample was taken)
ported by Divljan et al. (2006) who tested the method on were gained by plotting age in years against the natural log of
known-aged captive and wild grey-headed flying foxes the frequency of individuals in each age class (see Ricker 1975).
(Pteropus poliocephalus). The slope of the regression was taken as the instantaneous mor-
tality rate for all animals aged 2 years and above. Juveniles in
Determining the age of flying foxes the first year age category were assigned a mortality rate of 50%
Tooth preparation and sectioning owing to the inability to estimate a mortality rate for this age
class from the data. The natural survivorship function (lx) was
The two upper canine teeth, and their roots, were removed
calculated as:
intact from the jaw. The entire tooth above the gum line and at
the root base were cut off, leaving only the upper portion of the lx = N0(eMx)
root (Fig. 2). This made decalcification quicker and embedding
where Mx is the instantaneous mortality rate at time x and N0 is
of the tooth in wax easier, and ensured that all cross-sections
the number of animals at time 0. Annual rate of mortality (dx)
were taken from the same point of the root (the midpoint)
was estimated as:
because this part of the tooth is thought to provide the most con-
sistent approximation of age (Cool et al. 1994). dx = 1 eMx
Preparation for, and production of, tooth sections followed
Age at maturity and fecundity were calculated from the dataset
the protocol described in Cool et al. (1994), with some minor
in this study. Comparison of reproductive status data with age
adaptations. Sections cut at room temperature were difficult to
gave the proportion of females reproducing each year in this
obtain, so the exposed face of the tooth root was placed on tissue
sample (fecundity) and the age that females commenced pro-
paper and soaked in 10% ammonium hydroxide (on ice) for
ducing young. Fecundity (bx) is the number of female offspring
softening before sectioning. To stop sections lifting from slides
born per female per year and is calculated as:
during the staining process, we used poly-1-lysine slides with an
adhesive to maintain permanent contact between the section and bx = # of young born per season age-specific natality ratio
the slide. This was improved further by putting slides in a 37C of female to male offspring
oven for 3 days rather than the 60C for 24 h reported by Cool
The product of the survivorship function (lx) and fecundity (bx)
et al. (1994). The longer drying time allowed the sections to
gives average lifetime reproductive success (lxbx). The nett
adhere to the slides properly before staining.
reproductive rate R0 is calculated as:
Counting cementum layers R0 = ( lxbx)
We counted cementum layers on a double-headed Olympus
and is the number of viable female offspring produced by a
light microscope at 40 magnification. Several sections of each
single female in a lifetime. Mean generation time (G) is the
tooth were viewed to check for consistency in the number of
average age of mothers of new-born offspring and is calculated
cementum layers present. Counts were made in areas that exhib-
as:
ited clear layering across the entire width of the section. Any
sample for which it was difficult to assign the number of layers G = (lxbx x) /R0
was excluded from the analysis. As permanent dentition in
where x = the age of individuals. The intrinsic capacity for
flying foxes erupts soon after birth (Hall and Richards 2000), it
increase (rm) is an instantaneous rate and is derived from the
was not necessary to adjust the cementum layer counts (years)
generation time (G), which is only an estimate (Krebs 2001),
to allow for the number of years of life before the eruption of
permanent teeth.

Analysis
The frequency of individuals in each age class was plotted to
determine the shape of the age distribution for the overall popu-
lation, the shape for males and females separately, and a com-
parison of age distribution data for females between years. i.
Lactation data in females were graphed against age data to
determine when females first produced young, and the propor-
tion breeding each year.

Life tables
We constructed a life table for the Tolga Scrub colony fol-
lowing Krebs (2001) and Caughley (1977). As the fecundity of
females limits population growth, life tables calculating an
intrinsic capacity for increase use data on females only
(Caughley 1977; Stearns 1992; Caughley and Gunn 1996;
Krebs 2001). The maximum age (longevity) used was that of the
oldest individual aged in this study. Instantaneous mortality
ii.
iii.
Fig. 2. Diagram of tooth sectioning. (i) Above gum, exposed tooth,
removed before processing. (ii) Mid-section of root below gum-line,
removed to be embedded in wax. (iii) Basal section of root removed before
processing.
420 Wildlife Research
making rm also only an approximation when generations
overlap. A more accurate result is given with the finite rate of
increase (), following an iterative calculation using the
EulerLotka equation. The finite rate of increase () was calcu-
lated as:
= er
which is the exponential of r (or rm) (Krebs 2001). The age
distributions for 2001 and 2002 were compared using a
KolmagorovSmirnov two-sample test (Zar 1998) to determine
whether there was a significant difference between the years.
Life tables were completed for all females and pooled across
both years. Data for 2001 and 2002 were also separated and cal-
culated independently. To determine how vulnerable flying fox
populations are to differing rates of mortality and natality, we
conducted sensitivity analyses by creating additional life tables
with manipulated mortality and natality rates to estimate the
levels of these two factors needed to obtain a stable population.
Results
Age structure and demographic parameters
The age distribution of tick-paralysed P. conspicillatus indi-
viduals in 2001 and 2002 showed that 93% of the population
was 6 years of age or younger (Fig. 3). The maximum age was
13 years. The data for 2001 and 2002 were pooled once it was
determined that the age distributions for females for these
2 years were not significantly different (KolmagorovSmirnov
two-sample test, D = 0.23, P = 0.88).
Thirty-nine per cent of 2-year-old females had given birth
and by 3 years of age, most (80%) had reproduced (Fig. 4). Not
every female reproduced each year, with an average reproduc-
tive rate of 0.89 (89% of females producing a pup each year) for
females between 3 and 7 years of age. Reproductive rates for
females above 7 years were not calculated because sample sizes
were too small.
The instantaneous mortality rate was 43% for the popu-
lation overall and 40% for females alone. When this was con-
verted to an annual mortality rate, the overall rate was 35% and
33% for females. When survival was compared between males
and females with a log rank test, females had a significantly
greater survival rate than males (2358, 1 = 5.55, P < 0.05). The
90
80
Frequency of individuals
70
60
50
40
30
20
10
0
1 2 3 4 5 6
Age of Individuals in years
S. Fox et al.
overall sex ratio was 1 : 1 (2358, 1 = 2.52, P < 0.05, Yates cor-
rection applied).
Life-history tables
The net reproductive rate (R0) was estimated to be 0.364 per
generation (Table 1). The generation time (G) was ~5.0 years.
The intrinsic (or instantaneous) rate of population increase (rm)
was 18%. For this sample of the population during 20012002,
the more precise finite rate of increase () was 16.23%. When
the two years of data were treated independently, the finite rate
of increase () was 15.1% for 2001 and 12.01% for 2002.
Sensitivity analysis suggests that this sample population would
have been stable if the mortality rate was halved to 20%. If the
fecundity rate was increased to 100% for all females from the
age of 3 years, in addition to the reduced mortality rate, the
sample population would have increased by 1.6% per year.
Discussion
The analysis of population dynamics using life-history traits
and life tables is widely used in planning for the conservation
and management of wildlife (Stearns 1992; Saether et al. 1996;
Richard et al. 2002). For some animal groups, such as highly
mobile, inaccessible and nocturnal species like flying foxes,
acquiring the data needed for these analyses is extremely diffi-
cult (Pierson and Rainey 1992; Hall and Richards 2000; Vardon
and Tidemann 2000; McIllwee and Martin 2002; Barclay and
Harder 2003) and hence, this is the first field-databased life
table reported for a flying fox species.
Discussion of methods
Assumptions
The analysis assumes that the data collected are unbiased.
That is, individual animals have been sampled randomly
because the effect of the paralysis tick had no sex or age bias.
Several observations suggest this assumption is justified.
The sample is unlikely to be biased by age because the
overall age distribution graph (Fig. 3), and those of males and
females graphed separately (data not shown), suggest that all
age groups are exposed to tick paralysis. Also, the shape of the
age distribution curve (Fig. 3) is that expected of a mammal
with low reproductive output, high juvenile mortality and con-
Fig. 3. Age distribution for sample population
from Tolga Scrub colony on the Atherton Tablelands
for 20012002 (males and females combined).
Young-of-year of collection not included.
7 8 9 10 11 12 13
Indications of decline in the spectacled flying fox Wildlife Research 421

100

90
Percentage females lactating

80

70

60

50 Fig. 4. Proportion of females in each age class

found to be lactating. Sample sizes for age classes
40 8 years onward are low (8, n = 2; 9, n = 4; 11, n = 2;
30 13, n = 2).

20

10

0
1 2 3 4 5 6 7 8 9 10 11 12 13
Female age in years

stant mortality throughout the older age classes. In dogs, 80%. As this appeared excessive in the light of knowledge of the
younger dogs are more likely to survive tick paralysis than older birthing period and pup survival in the Tolga Scrub colony
dogs (Atwell et al. 2001), which suggests that older flying foxes (J. Maclean, unpubl. data), a more conservative juvenile mortal-
are not exempt from tick paralysis (or these data) by developing ity estimate of 50% was adopted. Vardon and Tidemann (2000)
an ability to resist the toxin. estimated that juvenile black flying foxes (P. alecto) have a mean
The sample is not biased to sex because the sex ratio of males first-year mortality rate of 68% and Sibly et al. (1997) found
to females in the sample was 1:1. Knowledge of sex ratios in juvenile mammals had a mean mortality rate of 56%, hence our
flying fox camps at different times of the year is limited but estimate of juvenile mortality could be considered conservative.
Nelson (1965) found sex ratios in P. poliocephalus nearing
equality at this time of the year. Does life history theory equate to actual traits?
The sample is also unlikely to be biased owing to sampling Although life-history theory is well understood, little is known
vagaries because all untreated animals envenomated by a paral- about whether life-history strategies, and their inherent trade-
ysis tick die (J. Maclean, unpubl. data), and in the 2 years that offs, still operate in species that are under threat. Spectacled
these data were collected every animal that fell in the focal flying foxes exhibit life-history traits expected of a species posi-
colony was collected. tioned at the slow end of the life history continuum. In the
Analysis using life tables assumes a closed population. This study sample, the age at first reproduction was 2 years, but only
assumption is problematic with mobile animals such as flying a minority of females gave birth at this age. It was not until
foxes but because of the timing of the study (the birthing season) females reached 4 years of age that they approached the mean
we believe it does not unduly affect the results. During the reproductive rate (89%) of mature females in the sample popu-
birthing period, mothers are tied to a particular colony by the
need to feed dependent young that are too heavy for the mother
to carry while foraging (Eby 1991). These young are left in Table 1. Life history table for female Pteropus conspicillatus
crches overnight while the mothers feed (Ratcliffe 1932) Symbols for columns: x, Age in years; lx, survivorship (mortality was set at
before returning each morning to the same colony and to their 50% for the first year of life and 40% for the remainder); bx, natality;
lxbx, age-specific reproductive rate; R0, net reproductive rate per generation;
waiting offspring. Mothers suckle their young until they are
G, mean generation time; r, intrinsic rate of increase of population
several months old, by which time the young are able to follow
adults to feeding trees at night (Eby 1991). As life-history tables x lx bx lxbx lxbxx R0 G r
are based solely on mortality and fecundity of adult females
(Krebs 2001), it is reasonable to consider the population as 0 1.000 0.000 0.000 0.000 0.364 5.049 0.1814
1 0.607 0.000 0.000 0.000
closed in the sampling period. Other assumptions made for this
2 0.407 0.000 0.000 0.000
analysis are that there is constant mortality within each age
3 0.273 0.395 0.108 0.323
group, that recruitment is constant for all age groups and that 4 0.183 0.455 0.083 0.332
there is a stable age distribution. These are standard assumptions 5 0.122 0.475 0.058 0.291
made in the use of life tables although it is recognised that they 6 0.082 0.445 0.037 0.219
are unlikely ever to be fully realised in nature (Caughley 1977; 7 0.055 0.500 0.028 0.193
Krebs 2001). 8 0.037 0.500 0.018 0.148
9 0.025 0.500 0.012 0.111
Juvenile mortality rate 10 0.017 0.500 0.008 0.083
Where the rate of juvenile mortality is unable to be mea- 11 0.011 0.500 0.006 0.061
12 0.007 0.500 0.004 0.045
sured, estimates of juvenile mortality are typically taken as
13 0.005 0.500 0.002 0.032
double the adult rate (Cailliet 1992), which in this case would be
422 Wildlife Research
lation. Delaying first breeding is common in species exhibiting
a slow life history, and although it might incur an immediate
cost to current reproductive output, it may also provide benefits
in terms of lifetime reproductive output or quality of offspring.
Delayed sexual maturity is usually offset by greater adult sur-
vival and the opportunity to reproduce many times in years to
come. This strategy results in more offspring being born to
bigger, healthier mothers with a greater probability of the off-
spring surviving (Stearns 1992). The reduction in overall repro-
ductive output from delayed age at first reproduction that this
entails, and the low overall reproductive rate, is believed to be a
significant predictor of extinction in the Megachiroptera (Jones
et al. 2003).
Most females (89%), from age 3 years onwards, produce one
young each year. This is consistent with studies on other species
of Pteropus (Vardon and Tidemann 1998; Barclay et al. 2004),
although Pteropus rodricensis was found to breed every alter-
nate birth cycle (West 1986). P. rodricensis is also critically
endangered
The generation time (G) was 5 years, suggesting that this is
the peak age of reproductive output in the colony. As few indi-
viduals live beyond the age of 6 years, there is little time
between the beginning of the major reproductive phase and the
probable end of life.
High mortality rate results in a lower-than-expected
longevity
The annual mortality rate of 35% for the sample population is
consistent with mortality data for mammals in general (Sibly
et al. 1997) but the combination of low reproductive output and
late age at first reproduction means that a stable population can
only be maintained if the majority of animals (especially
females) experience low year-to-year mortality. This was not the
case for our focal population of P. conspicillatus. The difference
between the annual mortality rate and the instantaneous mortal-
ity rate during 2001 and 2002 is likely to reflect the effect of tick
paralysis on the colony because both years were considered
bad tick paralysis seasons (J. Maclean, unpubl. data). Under
current circumstances of raised mortality (from ticks and other
causes), the usual trade-offs in this slow end species were not
operating effectively.
McIllwee and Martin (2002) suggested that Australian flying
foxes cannot sustain a mortality rate higher than 20%, irrespec-
tive of the rate of fecundity. The mortality rate for 2001 and
2002 in the Tolga Scrub colony exceeded this figure and
although it may be that losses to tick paralysis in those years
were unusually severe and the year-to-year mortality rate is
lower, calculations based on more usual levels of tick infestation
(J. Maclean, unpubl. data) still yield mortality rates in excess of
sustainable levels.
There is an extensive literature emphasising the long life
spans of bats (Read and Harvey 1989; Austad and Fischer 1991;
Pierson and Rainey 1992; Ransome 1995; Racey and Entwhistle
2000; Jonsson and Ebenman 2001; Barclay and Harder 2003;
Jones et al. 2003; Barclay et al. 2004; Brunet-Rossinni and
Austad 2004). In this study, few individuals reach senescence
(which in the spectacled flying fox exceeds 13 years of age) and
93% of animals investigated were 6 years of age or less. This
pattern has also been found in other bat species. Divljan et al.
S. Fox et al.
(2006) aged their oldest grey-headed flying fox at 23 years but
found that 85% of their sample of wild grey-headed flying foxes
were 6 years of age or less, and Ransome (1995) found that
although some female greater horseshoe bats (Rhinolophus
ferrumequinum) could still reproduce at the age of 29 years,
their average age at death was 8.1 years. This has obvious con-
sequences for demographic models because actual and expected
reproductive output will differ enormously, particularly where
captive animals are used as indicators of potential lifespan.
Life tables suggest a negative population growth rate
The life-history table analysis presented here suggests that, in
the absence of immigration, the spectacled flying fox colony at
Tolga Scrub declined by 16% as a result of high mortality rates
experienced during 2001 and 2002. Sampling was restricted to
a single colony and the analysis was conducted in the absence
of broader population data, so it is impossible to tell whether
this equates to a general decline across the species. It is worth
noting, however, that the Tolga Scrub colony is just one of
several colonies on the Atherton Tableland afflicted by similar
levels of tick envenomation and paralysis (J. Maclean unpub.
data). Immigration into the region exposes new animals to risk,
and the possibility of Tableland colonies acting as a population
sink (Hanski 1999) for the spectacled flying fox should not
be ignored.
Implications for management and conservation
Australian spectacled flying foxes suffer from many threats
(Richards and Hall 1998) and results from our study suggest that
the mortality rates experienced in 2001/2002 in the Tolga scrub
colony were twice as high as they could realistically sustain.
Elasticity analysis has shown that population growth rates of
mammals at the slow end of the life history continuum are
more sensitive to changes in adult and juvenile survival than
changes in reproductive parameters (Heppell et al. 2000;
Jonsson and Ebenman 2001; Oli 2004). This means conserva-
tion efforts directed towards preventing the decline of a
mammal species with a slow life history should focus on
trying to reduce mortality rates. As tick paralysis is capable of
exerting significant pressure on spectacled flying fox numbers,
especially on the Atherton Tablelands, efforts to reduce mortal-
ity through control of tick paralysis are required. Ticks are
picked up by spectacled flying foxes when they forage for fruit
on the low-lying introduced weed, Solanuum mauritianum,
which has become a common food source in the diet of specta-
cled flying foxes since the 1980s (Eggert 1994; Parsons et al.
2006). Eradication of this weed would not only reduce mortal-
ity rates in the Atherton Tablelands colonies, but also prevent
Solanuum mauritianum, and therefore tick paralysis, spreading
to the lowland colonies within the Wet Tropics region.
If the difference between the annual and instantaneous mor-
tality rates is removed (likely the effect of tick paralysis), the
mortality rate for the 2 years studied was still unsustainable.
This result suggests that tick paralysis, though a major addi-
tional risk to the spectacled flying fox, is only one of several
threats that place pressure on the Atherton Tablelands colonies.
Quantification of all threats faced by the spectacled flying fox is
an essential precondition to development of an effective recov-
ery or conservation plan for this species.
Indications of decline in the spectacled flying fox
Flying foxes around the world face continuing pressure for
space and food resources, as well as pressure from human
hunters. Although not considered a charismatic mammal group,
flying foxes have been widely recognised as ecologically and
economically important to the maintenance and continuation of
many endemic Pacific Island forests (Cox et al. 1991, 1992;
Wiles and Fujita 1992; McConkey and Drake 2006), where they
are often the only species large enough to disperse some seeds.
In Australia, flying foxes have been acknowledged as crucial in
pollination and seed dispersal of eucalypt forests and woodlands
(Eby et al. 1999; Palmer et al. 2000), and the spectacled flying
fox has been recognised as an important component of the Wet
Tropics World Heritage Area in north Queensland (Westcott
et al. 2001). Results from our study suggest that analyses of
mortality rates in other threatened flying fox species will be an
important step in understanding their sensitivity to disturbance
and that further erosion of populations could lead to loss of the
ecological services flying foxes provide, long before we recog-
nise them as being at risk (McConkey and Drake 2006).
Acknowledgements
We would like to thank the Rainforest Co-operative Research Centre,
Growcom, and Bat Conservation International for funding. SF was funded
by an ARC linkage grant. We thank Joanne Isaac, Tim Harvey, Ivan Lawler
and Euan Ritchie for comments on the manuscript, and Sue Reilly for help
in the Histology laboratory. This research is covered by ethics approval from
the James Cook University Animal Ethics sub-committee (# A723_02) and
Queensland Parks and Wildlife permits WISP00556402 and
WITK00554802.
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Manuscript received 3 September 2007, accepted 21 April 2008